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Garlic Mustard) on Morchella UNIVERSITY OF WISCONSIN-LA CROSSE Graduate Studies EFFECTS OF ALLIARIA PETIOLATA (GARLIC MUSTARD) ON MORCHELLA (MORELS) IN VITRO A Chapter Style Thesis Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science in Biology Elizabeth A. Leighton College of Science and Health Biology August, 2017 iii EFFECTS OF ALLIARIA PETIOLATA (GARLIC MUSTARD) LEACHATE ON MORCHELLA (MORELS) IN VITRO By Elizabeth Leighton We recommend acceptance of this thesis in partial fulfillment of the candidate’s requirements for the degree of Master of Science. The candidate has completed an oral defense of the thesis. __________________________________ _______________ Thomas Volk, Ph. D. Date Thesis Committee Chairperson __________________________________ _______________ Anita Baines, Ph. D. Date Thesis Committee Member __________________________________ _______________ Meredith Thomsen, Ph. D. Date Thesis Committee Member _______________________________ _______________ Todd Osmundson, Ph. D. Date Thesis Committee Member Thesis accepted ______________________________ ______________ Meredith Thomsen, Ph. D. Date Graduate Studies Director iv ABSTRACT Leighton, E. A. Effect of Alliaria petiolata (garlic mustard) on Morchella (morels) in vitro. MS in Biology, August 2017, 57pp. (T. Volk) Previous research has shown that Alliaria petiolata, garlic mustard, successfully invaded northern North America through allelopathy and suppressing arbuscular and ectomycorrhizal fungi. The research suggests that allelochemicals suppress mycorrhizal fungi’s spore viability and infectivity. Morels (Morchella) are popular edible fungi that live in habitats threatened by A. petiolata, even though their interaction with this plant has not been studied. This study explored whether Morchella is suppressed by A. petiolata by examining the effects of aqueous root, shoot, and whole plant A. petiolata leachates on Morchella cultures in vitro. Four traits appeared in response to treatment: deformation, line, pigment, and zone of inhibition. The root leachate elicited higher expression of pigment, line, and zone of inhibition traits. Morchella elata s.l. cultures were more affected by A. petiolata root leachate. Morchella americana cultures were likely to express line and pigment traits, but not the zone of inhibition trait. The in vitro conditions may not perfectly simulate soil conditions, but the responses in this experiment could occur in nature. Further research is needed to understand whether these traits appear in nature and whether these traits are associated with suppression in nature. iii ACKNOWLEDGMENTS I would like to thank Dr. Thomas Volk for all the patience, advice, and inspiration. This project would not have been possible without his guidance. Dr. Todd Osmundson has earned my gratitude for sound and calm advice in difficult times. Dr. Anita Baines was an enormous help with statistics and figures. Dr. Meredith Thomsen offered great insight and advice on experimental design. I also thank Dr. Bauman for being a great statistics resource. And thank you Beth Paluch for all the help, wisdom, and keeping the Petri dishes stocked. I also owe many thanks to my husband, Mark Schimpf, for the incredible amount of support and patience. He had so much great advice on graduate school and research. He was a rock for my entire graduate career. I would also like to thank my family for all the support they have given over the course of a lifetime. It’s far more than I can even measure. Lastly, I would like to thank those who sent black Morchella samples. JJ Murphy of New York, NY, Stef Lee of Wittenberg, WI, and Renée Lebeuf of Pierre Fonds, Quebec. This work would not have been possible without their generosity and help. iv TABLE OF CONTENTS PAGE LIST OF FIGURES ........................................................................................................... vi CHAPTER I: ALLIARIA PETIOLATA ALLELOPATHY AND INVASION IN NORTH AMERICA .......................................................................................................................... 1 Alliaria petiolata Invasion .............................................................................................. 1 Alliaria petiolata’s Allelochemistry................................................................................ 4 Allelopathy Case Studies .............................................................................................. 10 Research Objectives ...................................................................................................... 16 CHAPTER II: IN VITRO EXPOSURE OF MORCHELLA TO ALLIARIA PETIOLATA LEACHATE ..................................................................................................................... 18 Materials and Methods .................................................................................................. 22 Morchella Sampling and Culturing ........................................................................... 22 Table 1. Site data for isolates used in the study ................................................... 22 Leachate Preparation ................................................................................................. 23 Experimental Design ................................................................................................. 24 Statistical Analysis .................................................................................................... 26 Results ........................................................................................................................... 27 Deformation Trait ...................................................................................................... 28 Line Trait ................................................................................................................... 30 Pigment Trait ............................................................................................................. 31 Zone of Inhibition ...................................................................................................... 32 Isothiocyanate Control............................................................................................... 33 Discussion ..................................................................................................................... 35 Comparing Leachates ................................................................................................ 38 Exhibition of Traits by Clade .................................................................................... 40 Contextualizing Results ............................................................................................. 42 Future Experiments ................................................................................................... 43 Conclusion ................................................................................................................. 45 REFERENCES ................................................................................................................. 46 v LIST OF FIGURES FIGURE PAGE 1. Alliaria petiolata basal rosette and flowers.……………………………....................3 2. Chemical structures of sinigrin and allyl isothiocyanate………………….................5 3. Chemical structures of glucotropaeolin and benzyl isothiocyanate……….................5 4. Chemical structures of alliarinoside and isovitexin...………………………………..9 5. Morchella americana and M. angusticeps mushrooms…..…………...….................21 6. Diagram of methods………………………………………………………................24 7. Example of each trait observed…………………………………………...................26 8. The average proportion of positive responses for each trait and leachate…………..28 9. Proportion of positive deformation responses for all leachates in both black and yellow morels over time…………………………………………………..................29 10. Proportion of positive line responses for all leachates in both black and yellow morels over time……………………………………………………………………………..31 11. Proportion of positive pigment respoonses for all leachates in both black and yellow morels over time……………………………………………………………………..32 12. Proportion of positive zone of inhibition responses for all leachates in both black and yellow morels over time……………………………………………………………..33 13. Close-up image of deformation, line and pigment traits..…………………………...36 14. Example of the line and pigment traits………………………………………..……..42 vi CHAPTER I ALLIARIA PETIOLATA ALLELOPATHY AND INVASION IN NORTH AMERICA Alliaria petiolata Invasion Garlic mustard, Alliaria petiolata, is aggressively invading forest communities throughout much of North America. A great deal of research has been done to understand why it so successfully establishes large stands in North America when it does not do so in its native Europe. Studies have shown that the presence of A. petiolata in soil alters both the arbuscular and ectomycorrhizal fungi communities (Callaway et al., 2008; Stinson et al., 2006; Wolfe et al., 2008). However, not all mycorrhizal fungi have been studied; thus, our understanding of A. petiolata’s suppression is incomplete. To date, no published studies have reported on whether A. petiolata affects any edible mycorrhizal fungi. This study begins to explore whether A. petiolata affects the growth of Morchella, a highly prized edible mushroom. Rapid movement across the globe of many organisms has resulted in biological invasions in many environments worldwide. Invasive species are newly established in an environment and dominate
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