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Insights Into Triterpene Metabolism in Model Monocotyledonous and Oilseed Plants Genetically Engineered with Genes from Botryococcus Braunii

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Insights Into Triterpene Metabolism in Model Monocotyledonous and Oilseed Plants Genetically Engineered with Genes from Botryococcus Braunii University of Kentucky UKnowledge Theses and Dissertations--Plant and Soil Sciences Plant and Soil Sciences 2016 Insights into Triterpene Metabolism in Model Monocotyledonous and Oilseed Plants Genetically Engineered with Genes from Botryococcus Braunii Chase F. Kempinski University of Kentucky, [email protected] Author ORCID Identifier: http://orcid.org/0000-0002-5250-1291 Digital Object Identifier: https://doi.org/10.13023/ETD.2016.418 Right click to open a feedback form in a new tab to let us know how this document benefits ou.y Recommended Citation Kempinski, Chase F., "Insights into Triterpene Metabolism in Model Monocotyledonous and Oilseed Plants Genetically Engineered with Genes from Botryococcus Braunii" (2016). Theses and Dissertations--Plant and Soil Sciences. 80. https://uknowledge.uky.edu/pss_etds/80 This Doctoral Dissertation is brought to you for free and open access by the Plant and Soil Sciences at UKnowledge. It has been accepted for inclusion in Theses and Dissertations--Plant and Soil Sciences by an authorized administrator of UKnowledge. For more information, please contact [email protected]. STUDENT AGREEMENT: I represent that my thesis or dissertation and abstract are my original work. Proper attribution has been given to all outside sources. I understand that I am solely responsible for obtaining any needed copyright permissions. I have obtained needed written permission statement(s) from the owner(s) of each third-party copyrighted matter to be included in my work, allowing electronic distribution (if such use is not permitted by the fair use doctrine) which will be submitted to UKnowledge as Additional File. I hereby grant to The University of Kentucky and its agents the irrevocable, non-exclusive, and royalty-free license to archive and make accessible my work in whole or in part in all forms of media, now or hereafter known. I agree that the document mentioned above may be made available immediately for worldwide access unless an embargo applies. I retain all other ownership rights to the copyright of my work. I also retain the right to use in future works (such as articles or books) all or part of my work. I understand that I am free to register the copyright to my work. REVIEW, APPROVAL AND ACCEPTANCE The document mentioned above has been reviewed and accepted by the student’s advisor, on behalf of the advisory committee, and by the Director of Graduate Studies (DGS), on behalf of the program; we verify that this is the final, approved version of the student’s thesis including all changes required by the advisory committee. The undersigned agree to abide by the statements above. Chase F. Kempinski, Student Dr. Joe Chappell, Major Professor Dr. Arthur G. Hunt, Director of Graduate Studies INSIGHTS INTO TRITERPENE METABOLISM IN MODEL MONOCOTYLEDONOUS AND OILSEED PLANTS GENETICALLY ENGINEERED WITH GENES FROM BOTRYOCOCCUS BRAUNII DISSERTATION A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the College of Agriculture, Food and Environment at the University of Kentucky By Chase Foster Kempinski Lexington, Kentucky Director: Dr. Joe Chappell, Professor of Plant Physiology Lexington, Kentucky 2016 Copyright © Chase Foster Kempinski 2016 ABSTRACT OF DISSERTATION INSIGHTS INTO TRITERPENE METABOLISM IN MODEL MONOCOTYLEDONOUS AND OILSEED PLANTS GENETICALLY ENGINEERED WITH GENES FROM BOTRYOCOCCUS BRAUNII Isoprenoids are one of the most diverse classes of natural products and are all derived from universal five carbon, prenyl precursors. Squalene and botryococcene are linear, hydrocarbon triterpenes (thirty carbon compounds with six prenyl units) that have industrial and medicinal values. Squalene is produced by all eukaryotes as it is the first committed precursor to sterols, while botryococcene is uniquely produced by the green algae, Botryococcus braunii (race B). Natural sources for these compounds exist, but there is a desire for more renewable production platforms. The model plant Arabidopsis thaliana was engineered to accumulate botryococcene and squalene in its oil seeds using a combination of biosynthetic genes and subcellular targeting strategies. The model monocotyledonous grass, Brachypodium distachyon, was also engineered to accumulate botryococcene in its leaves, testing a variety of subcellular targeting methods. Both oilseeds and grasses have highly desirable characteristics as production platforms, and both can utilize photosynthesis to power the biosynthesis of these valued hydrocarbons. In each of these efforts, we were able to obtain high levels of triterpene accumulation and uncovered new aspects of isoprenoid metabolism and its regulation. Also investigated was a new gene from the green algae, Botryococcus braunii (race B), encoding for a novel methyltransferase, which in combination with a previously reported methyltransferase, was characterized as converting dimethylated squalene to tetramethylated. Tetramethylated derivatives of squalene (and botryococcene) are highly desirable for industrial applications, and the discovery of the genes encoding for this biosynthetic capacity portends opportunities to engineer other heterologous host organisms and create other amenable production platforms. KEYWORDS: isoprenoid, triterpene, genetic engineering, Brachypodium, Arabidopsis, Botryococcus braunii Chase Foster Kempinski Student’s Signature 09-30-2016 Date INSIGHTS INTO TRITERPENE METABOLISM IN MODEL MONOCOTYLEDONOUS AND OILSEED PLANTS GENETICALLY ENGINEERED WITH GENES FROM BOTRYOCOCCUS BRAUNII By Chase Foster Kempinski Joe Chappell Director of Dissertation Arthur G. Hunt Director of Graduate Studies 09-30-2016 ACKNOWLEDGEMENTS All of the work presented here would not have been possible without the support, encouragement, and freedom to explore given by my adviser, Joe Chappell. His confidence in me allowed me to push through the setbacks and persevere, learning that science is as much about persistence as it is about inspiration. I feel that this dissertation is only a small reflection of all the knowledge and skills that I have obtained during my time in his laboratory, and for that I thank him. I also want to thank my committee members, Arthur Hunt, Sharyn Perry, and Seth DeBolt for their time, comments, critiques and encouragement during my work. I also thank my outside examiner, Edgar Cahoon, for his donation of materials for some of the projects here and for his time and critical comments of this dissertation. I want to acknowledge all of Chappell lab members that have helped me along my path, especially Scott Kinison, Zuodong Jiang, and Stephen Bell. Their support and conversations helped me get started and maintain a good course during my time here. Importantly, I wish to thank my family, because without their support through all my collegiate career, none of this would be possible. Everything that I have been able to accomplish I owe to them. iii TABLE OF CONTENTS ACKNOWLEDGEMENTS ............................................................................................... iii TABLE OF CONTENTS ................................................................................................. iv LIST OF TABLES .......................................................................................................... vii LIST OF FIGURES ....................................................................................................... viii CHAPTER 1: ENGINEERING ISOPRENOID METABOLISM IN PLANTS—A PRIMER . 1 1.1 Introduction ........................................................................................................... 1 1.2 Host choice ........................................................................................................... 5 1.3 Organ and tissue-specific engineering .................................................................. 7 1.3.1 Leaves ............................................................................................................ 8 1.3.2 Seeds ............................................................................................................10 1.3.3 Fruits .............................................................................................................12 1.3.4 Roots .............................................................................................................13 1.3.5 Other specialized cell types ...........................................................................15 1.4 Intracellular pathway engineering .........................................................................16 1.5 Understanding and utilizing compartmentalization ...............................................20 1.6 Conclusions and future directions ........................................................................22 CHAPTER 2: ENGINEERING TRITERPENE METABOLISM IN THE MODEL MONOCOT, BRACHYPODIUM DISTACHYON .................................................................................28 2.1 Summary .............................................................................................................28 2.2 Introduction ..........................................................................................................28 2.3 Results .................................................................................................................32 2.3.1 Elevated levels of cytosolic botryococcene are dependent on the inclusion of a FPS gene in combination with BS ...........................................................................32
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