Florida State University Libraries Electronic Theses, Treatises and Dissertations The Graduate School 2013 Evolutionary Constraints on Plasticity in the Anti-Herbivore Defenses of Solanum Carolinense David William McNutt 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 EVOLUTIONARY CONSTRAINTS ON PLASTICITY IN THE ANTI-HERBIVORE DEFENSES OF SOLANUM CAROLINENSE By DAVID WILLIAM MCNUTT A Dissertation submitted to the Department of Biological Science in partial fulfillment of the requirements for the degree of Doctor of Philosophy Degree Awarded: Fall Semester, 2013 David W. McNutt defended this dissertation on October 15, 2013. The members of the supervisory committee were: Nora Underwood Professor Directing Dissertation Frederick Davis University Representative Brian Inouye Committee Member Joseph Travis Committee Member Alice Winn Committee Member The Graduate School has verified and approved the above-named committee members, and certifies that the dissertation has been approved in accordance with university requirements. ii ACKNOWLEDGMENTS Many thanks to my doctoral committee for their helpful comments and advice during my graduate career and the preparation of this dissertation. I thank Nora Underwood for being a great advisor in every sense of the word. She has provided invaluable advice regarding my research and professional development, and served as an excellent faculty role model. I thank Brian Inouye for putting up with my unending statistics questions and Alice Winn for cultivating my teaching abilities and the many conversations that helped me wrap my head around phenotypic plasticity. I also thank Joe Travis for all the enjoyable Friday afternoons; his paper discussion groups were stimulating and exposed me to many new ideas. This work would have not been possible without the assistance of countless volunteers and field assistants. I am particularly indebted to Amanda Buchanan for her field/greenhouse help and the many intellectual discussions that strengthened my dissertation. I also thank the FSU greenhouse staff (Karen Graffius and Theresa Jepsen) and the NFREC field crew (Tom Bolton) for logistical help with experiments, Stacey Halpern for providing me with Solanum clonal lines, and Tania Kim for being so generous when dividing roots among our various projects. I thank Megan Mayo, Ian Padykula, Kylie Rafferty, Ryan Coker, and Brittany Forer not only for the many hours they spent surveying and processing plants, but for entertaining me in the lab and field. And of course, I thank all of the past and current members of the Inouye- Underwood labs and Muffins for Mom for keeping me sane. Parts of this dissertation were funded by a Robert K. Godfrey Endowment in Botany, an FSU Planning Grant, an NSF Doctoral Dissertation Improvement grant, and two grants to Nora Underwood: NSF DEB-0717221 and USDA grant #2005-35302-16311. iii TABLE OF CONTENTS List of Tables ................................................................................................................................. vi List of Figures ............................................................................................................................... vii Abstract ........................................................................................................................................ viii 1. INTRODUCTION ...................................................................................................................1 2. RESISTANCE TRADE-OFFS ACROSS PLANT GENOTYPES STRUCTURE INDIRECT INTERACTIONS AMONG HERBIVORES......................................................4 2.1 Introduction ....................................................................................................................4 2.2 Methods..........................................................................................................................7 2.1.1 Study Species .....................................................................................................7 2.2.2 Experimental Methods .......................................................................................8 2.2.3 Statistical Analyses ............................................................................................9 2.3 Results ..........................................................................................................................12 2.4 Discussion ....................................................................................................................17 3. GENETIC CONSTRAINTS ON THE EVOLUTION OF INDUCED ANTI-HERBIVORE DEFENSES AND TOLERANCE ........................................................................................22 3.1 Introduction ..................................................................................................................22 3.2 Methods........................................................................................................................26 3.2.1 Study System ...................................................................................................26 3.2.2 Experimental Methods .....................................................................................27 3.2.3 Trait Measurements .........................................................................................28 3.2.4 Vegetative Community Metrics .......................................................................31 3.2.5 Statistical Analyses ..........................................................................................31 3.3 Results ..........................................................................................................................34 3.4 Discussion ....................................................................................................................37 4. THE ADAPTIVE VALUE AND COSTS OF PLASTICITY IN PLANT INDUCED DEFENSES AND TOLERANCE OF HERBIVORY ..........................................................43 4.1 Introduction ..................................................................................................................43 4.2 Methods........................................................................................................................47 4.2.1 Study System and Field site .............................................................................47 4.2.2 Experimental Methods .....................................................................................48 4.2.3 Trait Measurements .........................................................................................49 4.2.4 Statistical Analyses ..........................................................................................53 iv 4.3 Results ..........................................................................................................................58 4.4 Discussion ....................................................................................................................65 5. CONCLUSIONS ...................................................................................................................73 APPENDICES ...............................................................................................................................77 A. SUPPLEMENTARY MATERIALS FOR CHAPTER TWO ...............................................77 B. SUPPLEMENTARY MATERIALS FOR CHAPTER THREE ...........................................79 C. SUPPLEMENTARY MATERIALS FOR CHAPTER FOUR .............................................84 REFERENCES ..............................................................................................................................89 BIOGRAPHICAL SKETCH .......................................................................................................104 v LIST OF TABLES 3.1 Defense traits measured in this study and evidence for their defensive value and damaged-induced plasticity ...............................................................................................26 4.1 Adaptive value of plasticity analysis .................................................................................60 4.2 Costs of tolerance ...............................................................................................................65 A.1 Results of three-way, fixed factor ANOVAs testing for effects of block, genotype, and damage treatment on RGR of L. juncta larvae after 48hrs of feeding ...............................77 A.2 Results of three-way, fixed factor ANOVAs testing for effects of block, genotype, and damage treatment on RGR of M. sexta larvae after 48hrs of feeding ................................78 A.3 Pearson correlation coefficients between genotypic measures of induced and constitutive resistance to L. juncta (LJ) and M. sexta (MS). .................................................................78 B.1 Genetic variation in the mean expression of defense traits ................................................80 B.2 Genetic variation in the plasticity of defense traits ............................................................81 B.3 Genetic variation in tolerance ability. ................................................................................81 B.4 Genetic correlations among the constitutive expression of five defense traits and three measures of tolerance. ........................................................................................................82