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Information to Users INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6” x 9” black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. UMI A Bell & Howell Infonnation CompaiQr 300 Noith Zed) Road, Ann Aibor MI 48106-1346 USA 313/761-4700 800/521-0600 INVESTIGATION OF TUBULINS IN ASPERGILLUS NIDUIANS AND CYANIDIUM CALDARIUM DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Yassmine M. Nazih Akkari, B. S. ***** The Ohio State University 1997 Dissertation Committee: Dr. Berl R. Oakley, Adviser Dr. Thomas J. Byers pproved^y Dr. George A. Marzluf Dr. Amanda A. Simcox Dr. Dale D. Vandre Adviser Department of Molecular Genetics UMI Number: 9721068 UMI Microform 9721068 Copyright 1997, by UMI Company. All rights reserved. This microform edition Is protected against unauthorized copying under Title 17, United States Code. UMI 300 North Zeeb Road Ann Arbor, MI 48103 ABSTRACT Microtubules are essential for a variety of functions in eukaryotic cells. They are formed from molecules of tubulin, a heterodimer consisting of two globular proteins called a and P tubulin. A third member of the tubulin superfamily, y tubulin, is located at the microtubule organizing centers and is required for microtubule assembly in vivo. Surprisingly, little is known about the details of microtubule structure and assembly. The determination of the three-dimensional structure of tubulin, can greatly contribute to a better understanding of microtubule assembly at the molecular level. However, progress in the elucidation of the tubulin tertiary structure has been hindered by the heterogeneity of tubulin and its intrinsic lability. To address these issues, I have puriried assembly-competent and homogeneous tubulin from Aspergillus nidulans and investigated the tubulin genes from the thermophilic alga Cyanidiwn caldarium as a potential source of thermostable tubulins. A. nidulans has two ^-tubulin genes {benA and tubC) and two a-tubulin genes (tubA and tubB). In order to isolate homogeneous tubulin, I have constructed a strain with only one functional a-tubulin gene, tub A, and in which six histidines have been inserted at the C-terminus of the benA gene. This allowed purifrcadon of tubulin using a nickel column. I have used electron microscopy and two-dimensional gel electrophoresis to show that this purified tubulin is assembly-competent and isotypically pure. In addition, since thermophilic organisms are a potential source of heat stable proteins, and tubulins from A. nidulans may still prove to be labile, I have investigated the tubulin genes from the thermophilic red alga, Cyanidium caldarium . I have sequenced internal fragments of genes encoding one a tubulin, four p tubulins and one y tubulin from C. caldarium, generated by PCR. Subsequently, I have constructed a C. caldarium genomic library, cloned the full-length copies of the four p-tubulin genes and the y-tubulin gene, and sequenced the y-tubulin gene and P 1-tubulin gene. I have also investigated the total number of tubulin genes from this organism and the genomic arrangement of its p-tubulin genes. m Dedicated to my father Nazih Akkari and my mother Nada Zock IV ACKNOWLEDGMENTS I wish to thank my advisw. Dr. Berl R. Oakley, for his intellectual support and encouragement, his great scientific insight, and for giving me the opportunity to work on different projects and learn different techniques. He has taught me the value of a careful and rigorous scientific training and for this, I will always be grateful. I also wish to thank my committee members, Drs. Thomas J. Byers, George A. Marzluf, Amanda A. Simcox, and Dale D. Vandre for their time and helpful suggestions. I would especially like to thank Dr. Tom Byers for his encouragement, and Mandy Simcox for her friendship and continued interest in my career. I would like to thank Dr. Bill Birky for helpful discussions and Dr. Paul Fuerst, Sulayman Dib-Hajj and 2^eina Daoud for helping me to fulfill my dream of pursuing a graduate career in the United States. In the Oakley lab, I would like to thank Kathy Jung for the generosi^ of her heart, and for being a great friend and mentor, and Liz Oakley and Dawnne Wise for their support and friendship. I would also like to thank previous members of our lab: Mary Ann Martin, Yisang Yoon, Tetsuya Horio and Tomohiro Akashi for their friendship and helpful discussions. In addition, I would like to thank Gary Grumbling for his friendship, his expert computer assistance, and his help with the C. caldarium tubulin sequence analysis. I also wish to thank Don Ordaz for his help in growing A. nidulans cultures, and Jose Diaz and Kathy Wolken for their assistance in the use of the electron microscope. Most importantly, I would like to express my deepest love and gratitude to all the members of my family in Lebanon. To my parents, I owe my most sincere thanks and appreciation for their love, emotional and intellectual support, and for teaching me the good value of education and integrity in a time and place where these words seemed to have lost their meaning. They have always provided me with positive energy to reach my goals. I would also like to thank my sisters, Nesrine and Nada, and the rest of my family, especially my uncles Nabil Akkari and Aref Zock, for their love and faith in me. To my dearest Mend, Jeff Seiple, I express my sincere thanks for his love, for accepting my weaknesses and reinforcing my strengths, and for everything that he is. I am truly blessed to have found a person like him, and I will always be grateful for the happiness he brings to my life. I would also like to thank Jeffs family for their warmth and generosity and for considering me a part of their family. To all my wonderful friends, both in Lebanon and in the United States, I express my thanks for their unconditional love and support. I would especially like to thank my best friends Zeina Daoud, Hanady Salman and Ghina Dabboussy for the goodness of their hearts, their love and wonderful companionship. vi WA January 22,1969 Bora - Tripoli, Lebanon 1989................................................... B. S. Biology, American Univerrity of Beirut, Beirut, Lebanon 1990-presenL ......................................Graduate Teaching and Research Associate, Department of Molecular Genetics, The Ohio State University, Columbus, Ohio PUBUCATIONS 1. Y. Akkari, R. G. Bums, C. E. Oakley, and B. R. Oakley. 1994. Cloning and sequencing of y- and P-tubulin genes from the thermophilic alga Cyanidium calc^rium. Molec. Biol. Cell 55:282a. 2. Y. Akkari, Y. Yoon, and B. R. Oakley. 1995. Expression and purifrcation of isotypically pure tubulin from Aspergillus nidulans. Molec. Biol. Cell 55:32a. FIELDS OF STUDY Major Field: Molecular Genetics YU TABLE OF CONTENTS EagÊ Abstract ......................................................................................................... ii Dedication ..................................................................................................... iv Acknowledgments ........................................................................................ v Vita................................................................................................................. vii List of Tables .................................................................................................. xi List of Figures ................................................................................................ xii Chapters: 1. INTRODUCTION.............................................................................. 1 Overview ................................................................................. 1 General structure of microtubules .......................................... 2 Advances in the elucidation of microtubule structure 5 In vitro microtubule assembly and dynamics .......................... 6 Microtubule organizing centers and microtubule polarity .............................................. 13 Microtubule associated proteins ............................................ 17 Tubulin biochemistry: domain structure and drug interactions .................................................... 18 Tubulin molecular biology and genetics ................................ 22 Y tubulin ..................................................................................
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