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Distribution Agreement in Presenting This Thesis As a Partial Fulfillment Of Distribution Agreement In presenting this thesis as a partial fulfillment of the requirements for a degree from Emory University, I hereby grant to Emory University and its agents the non-exclusive license to archive, make accessible, and display my thesis in whole or in part in all forms of media, now or hereafter now, including display on the World Wide Web. I understand that I may select some access restrictions as part of the online submission of this thesis. I retain all ownership rights to the copyright of the thesis. I also retain the right to use in future works (such as articles or books) all or part of this thesis. Kevin Hoang April 15, 2016 Effects of milkweed species on monarch butterflies and their parasites by Kevin Hoang Jacobus de Roode Adviser Department of Biology Jacobus de Roode Adviser Levi Morran Committee Member Matthew Weinschenk Committee Member 2016 Effects of milkweed species on monarch butterflies and their parasites By Kevin Hoang Jacobus de Roode Adviser An abstract of a thesis submitted to the Faculty of Emory College of Arts and Sciences of Emory University in partial fulfillment of the requirements of the degree of Bachelor of Sciences with Honors Department of Biology 2016 Abstract Effects of milkweed species on monarch butterflies and their parasites By Kevin Hoang Parasites contribute to much of the species diversity and cover a wide range of hosts and environments. By definition, parasites induce fitness reductions in hosts, whether it is through reduction in host size, immune defense, lifespan, or fecundity. The intricate host-parasite relationship is affected by a number of factors, such as host diet, nutrition, and immune system. For example, the monarch butterfly (Danaus plexippus), which can be infected with the protozoan parasite Ophryocystis elektroscirrha, exhibits oviposition preferences when infected. Infected females, which spread the parasite during oviposition, seek out medicinal milkweeds (Asclepias spp.) to lay their eggs upon in order to alleviate the effects of O. elektroscirrha in the next generation. These medicinal milkweeds differ in their toxic chemical contents and reduce parasite loads as well as increase lifespans of infected monarchs compared to those reared on non-medicinal milkweeds. Here, we explore how different milkweed species affect the host- parasite interaction between the monarch butterfly and its parasite. Specifically, we compare Morrenia odorata, a member of the milkweed family, as a host plant for monarchs with more common milkweed species (Asclepias spp.). We found that M. odorata can be used to rear monarch larvae to adulthood but that the lifespans of these monarchs are generally lower than those reared on Asclepias milkweeds. Additionally, M. odorata does not appear to have any medicinal effects on infected monarchs. On top of this, female monarchs significantly preferred to oviposit on Asclepias milkweeds over M. odorata when given a choice. A separate experiment we conducted was to study the effects of different monarch larval diets on parasite morphology. Here, we found a significant relationship between larval diet and parasite size. The toxic chemicals within milkweeds significantly reduced parasite size, a hidden benefit of medicinal milkweeds on O. elektroscirrha. Not only do these milkweeds reduce parasite loads, but they also reduce parasite sizes, another possible mechanism to lower parasite fitness. In the future, additional studies on the effects of M. odorata and monarch larvae diets on monarch-parasite interactions need to be conducted to further expand our knowledge on this intricate and alluring host-parasite relationship. Effects of milkweed species on monarch butterflies and their parasites By Kevin Hoang Jacobus de Roode Adviser A thesis submitted to the Faculty of Emory College of Arts and Sciences of Emory University in partial fulfillment of the requirements of the degree of Bachelor of Sciences with Honors Department of Biology 2016 Acknowledgements I am very grateful to have had the opportunity to study in the de Roode lab. Dr. Jacobus de Roode has taught me so much in the past few years and has been a role model for all of the lab members to follow. I especially want to thank Dr. Leiling Tao for being my direct mentor and for giving me quick feedback throughout these projects. I want to thank previous and current de Roode lab members for teaching me how to and for helping rear caterpillars and milkweed plants as well as for their feedback and support during the projects. I am also very grateful for having Dr. Matthew Weinschenk and Dr. Levi Morran on my committee. Dr. Weinschenk's passionate lecture styles and mentorship inspired me to be an organic chemistry mentor, an experience that has helped me in presenting and explaining ideas and concepts to others. Dr. Morran always offers excellent advice and taking his course exposed me to scientific literature on experimental evolution. Finally, I would like to thank my professors here at Emory University for guiding me to be the student, researcher, and person that I am today. TABLE OF CONTENTS Chapter 1 Effects of Morrenia odorata on monarch fitness and parasite infection ABSTRACT .....................................................................................................................................1 INTRODUCTION ...........................................................................................................................1 MATERIALS AND METHODS .....................................................................................................5 Host and Parasite sources.....................................................................................................5 Plant sources ........................................................................................................................6 Experiment 1 ........................................................................................................................6 Experiment 2 ........................................................................................................................9 Statistical tests ......................................................................................................................9 RESULTS ......................................................................................................................................10 Experiment 1 ......................................................................................................................10 Summer 2015 .........................................................................................................10 Fall 2015 ................................................................................................................11 Experiment 2 ......................................................................................................................11 DISCUSSION ................................................................................................................................11 FIGURES .......................................................................................................................................14 Chapter 2 Host diet affects the morphology of a butterfly pathogen ABSTRACT ...................................................................................................................................22 INTRODUCTION .........................................................................................................................22 MATERIALS AND METHODS ...................................................................................................26 Host and Parasite sources...................................................................................................26 Plant sources ......................................................................................................................26 Experimental design...........................................................................................................27 Parasite morphology analysis ............................................................................................29 Statistical tests ....................................................................................................................29 RESULTS ......................................................................................................................................30 DISCUSSION ................................................................................................................................31 REFERENCES ..............................................................................................................................34 TABLES ........................................................................................................................................40 FIGURES .......................................................................................................................................43 TABLES AND FIGURES Chapter 1 Effects of Morrenia odorata on monarch fitness and parasite infection FIGURES Fig. 1 A general structure of cardenolides ...........................................................................3 Fig. 2 Percent of monarch survival to adulthood (Summer 2015) ....................................14 Fig. 3 Monarch lifespans by plant species (Summer 2015) ...............................................15 Fig. 4 The infection probability of monarchs reared on different milkweed species
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