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The Effects of Dietary Nutrient Balance on Life-History Traits and Sexual Selection in the Field Cricket, Gryllus Veletis

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The Effects of Dietary Nutrient Balance on Life-History Traits and Sexual Selection in the Field Cricket, Gryllus Veletis The effects of dietary nutrient balance on life-history traits and sexual selection in the field cricket, Gryllus veletis by Sarah J. Harrison A thesis submitted to the Faculty of Graduate and Postdoctoral Affairs in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biology Carleton University Ottawa, Ontario © 2018 Sarah J. Harrison Abstract Nutrition is an important driver of biological variation. Macronutrients such as protein and carbohydrates, and elemental nutrients such as phosphorus, are known to affect animal fitness traits. No study, however, has investigated the importance of phosphorus relative to dietary protein or carbohydrates, or their interactive effects, on animal performance. To advance our understanding of the impact of nutrition on individual fitness, my thesis examined the influence of dietary protein, carbohydrate, and phosphorus balance on fitness-related life-history traits, including those involved in intra- and inter-sexual selection, of Gryllus veletis field crickets. My findings revealed that adult lifespan, weight gain, males’ acoustic mate attraction signals, and females’ egg production were maximized on diets with different protein:carbohydrate ratios, such that not all fitness traits could simultaneously be maximized on the same diet. Similarly, juvenile females could not simultaneously maximize their growth, development rate, survival, and dispersal capability at adulthood on the same dietary protein:carbohydrate ratio. Adult males and females also had different optimal nutrient intake ratios for reproductive performance. My results support theoretical predictions for the condition- dependence of traits involved in inter- and intra-sexual selection; both male mate attraction signals, and female sexual responsiveness and preferences for such signals, were influenced by dietary protein:carbohydrate ratio. However, male aggressiveness in agonistic contests with rivals was not influenced by dietary nutrient balance. Contrary to my expectations, dietary phosphorus had little influence on fitness traits, with the exception of a negative influence of high phosphorus diets on male mate attraction ii signals. When given a choice between diets containing differing but complementary nutrient compositions, both adults and juveniles demonstrated an ability to regulate their protein and carbohydrate intakes, but not their phosphorus intake. These self-selected diets often represented a compromise between the differing nutrient requirements of multiple fitness traits. Overall, my findings suggest that environmental heterogeneity in nutrient availability is an important driver of variation in animal fitness, and highlight the importance of disentangling the influences of different nutrients on animal fitness traits. iii Preface Thesis format and statement of co-authorship: This dissertation is written as an ‘integrated thesis’ and contains four data chapters and a General Methods chapter, ‘bookended’ by a General Introduction chapter and a General Conclusions chapter. The data chapters are written in manuscript format and, for this reason, there is some repetition across chapters in their introduction and discussion sections that was necessary to properly frame my research within the existing scientific literature. Two data chapters (3 & 4) have been published, and the remaining chapters (5 & 6) are ready to be submitted for publication. While the research contained within this thesis is entirely my own, several co-authors contributed to the experimental designs and the writing of manuscripts. Below I list the contributions of myself and any co-authors for each data chapter. Chapter 3: Influence of dietary nutrient balance on male and female lifespan, growth, egg production and acoustic mate attraction signals This chapter is published as: Harrison, S. J., Raubenheimer, D., Simpson, S. J., Godin, J.-G. J., & Bertram, S. M. (2014). Towards a synthesis of frameworks in nutritional ecology: interacting effects of protein, carbohydrate and phosphorus on field cricket fitness. Proceedings of the Royal Society B: Biological Sciences, 281, 20140539. For this chapter, my co-supervisors, Drs. Susan M. Bertram and Jean-Guy J. Godin, and I collaboratively developed the research questions and designed the experiments. Drs. D. iv Raubenheimer and S.J. Simpson assisted with the diet treatment aspect of experimental design and advised on data analysis. I collected the data, performed the data analysis, and wrote the first draft of the manuscript. Dr. Raubenheimer assisted with writing the Introduction in the final draft of the manuscript. All co-authors provided comments and editorial suggestions which greatly improved the manuscript. Chapter 4: The influence of dietary nutrient balance on male aggression and signalling This chapter is published as: Harrison, S. J., Godin, J.-G. J., & Bertram, S. M. (2017). Influence of dietary nutrient balance on aggression and signalling in male field crickets. Animal Behaviour, 134, 123–134. For this chapter, Dr. Bertram, Dr. Godin, and I collaboratively developed the research question and designed the experiment. I collected the data, performed the data analysis, and wrote the manuscript. Drs. Bertram and Godin provided comments and editorial suggestions which greatly improved the manuscript. Chapter 5: The influence of dietary nutrient balance on female life-history traits throughout development and adulthood and v Chapter 6: The influence of dietary nutrient balance on female sexual responsiveness and mate preferences For these chapters, Dr. Bertram, Dr. Godin, and I collaboratively developed the research questions and designed the experiments. I collected the data, performed the data analyses, and wrote the manuscripts. Drs. Bertram and Godin provided comments and editorial suggestions which greatly improved these manuscripts. Permissions to include previously published, co-authored materials in this thesis: I have obtained permission from all co-authors to use our collaborative published papers in this thesis. Additionally, I have obtained permission from Royal Society Publishing and Elsevier Publishing to use the published papers associated with Chapters 3 and 4 in this thesis. To cite this thesis, I recommend: Harrison, S.J. 2018. The effects of dietary nutrient balance on life-history traits and sexual selection in the field cricket, Gryllus veletis. Ph.D. thesis, Carleton University, Ottawa, Ontario, Canada. vi Acknowledgements I am immensely grateful to my supervisors, Drs. Susan Bertram and Jean-Guy Godin, for their patience, encouragement, support and guidance over the past eight years. I thank them for their enormous contribution to my research and the completion of this thesis, and especially for putting up with my writing at a ‘snail’s pace’. Without their mentorship, I would not be the scientist I am today. A special thanks to Sue for twisting my arm into doing a Master’s degree, and then when I was almost free, convincing me to stay to do a Ph.D. I am also grateful to Carleton University staff and faculty for their support during my doctoral studies. I would especially like to thank the Biology Department’s amazing support staff for their ability to fix all of my administrative and greenhouse problems. I would also like to thank Drs. Howard Rundle and Jeff Dawson for serving on my thesis advisory committee, and for their thought-provoking questions, helpful advice, and encouragement over these past few years. A special thanks to Dr. Dawson for frequently helping me with experimental equipment repairs, design, and construction. Also a big thank you to all of the Bertram lab members both past and present for their assistance with animal care and experimental brainstorming, including Genevieve Ferguson, Mykell Reifer, Emily McAuley, Amy Villarreal, Michelle Loranger, Ian Thomson, Karen Pacheco, Caitlin Grant, Lauren Fitzsimmons, as well as countless undergraduate students. Likewise, thanks to my ‘half lab mates’ in the Godin lab. A special thanks to my friend and ‘lab wife’ Genevieve, my shoulder to complain on and vii partner in procrastination, without whom these past few years would have been significantly less entertaining and awesome. The completion of this doctoral thesis would not have been possible without the support of my family. An enormous thank you to my partner, Joe Chaput, for being an unending source of support, for always being there for me, and not letting me give up when finishing this thesis seemed too far from my grasp. Thanks to my parents, Bev and Maria, for providing me with the guidance, encouragement, and opportunities in life that got me to where I am today. I am especially grateful for their combined love of science and nature, which most definitely inspired my own love of biology. Funding for the research presented in this thesis was provided through research grants awarded to my supervisors, Drs. Susan Bertram and Jean-Guy Godin, from the Natural Sciences and Engineering Research Council (NSERC) of Canada Discovery Grants, the Canadian Foundation for Innovation, the Ontario Research Fund, and the Carleton University Research Fund. I was directly supported by Departmental Scholarships from Carleton University, an NSERC Canadian Graduate Scholarship, and an Ontario Graduate Scholarship. Lastly, I would also thank the journal editors and anonymous reviewers for their extensive helpful comments on the published manuscripts which were based on Chapters 3 and 4. viii Table of Contents Abstract ..............................................................................................................................
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