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Taxonomy, Molecular Phylogeny and Taxol Production in Selected Genera of Endophytic Fungi by Jeerapun Worapong a Dissertation Su

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Taxonomy, Molecular Phylogeny and Taxol Production in Selected Genera of Endophytic Fungi by Jeerapun Worapong a Dissertation Su Taxonomy, molecular phylogeny and taxol production in selected genera of endophytic fungi by Jeerapun Worapong A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Plant Pathology Montana State University © Copyright by Jeerapun Worapong (2001) Abstract: This study examined the taxonomy, molecular phytogeny, and taxol production in selected genera of endophytic fungi associated with tropical and temperate plants. These common anamorphic endophytes are Pestalotiopsis, Pestalotia, Monochaetia, Seiridium, and Truncatella, forming appendaged conidia in acervuli. Sexual states of these fungi, including Amphisphaeria, Pestalosphaeria, Discostroma and Lepteutypa, are in a little known family Amphisphaeriaceae, an uncertain order of Xylariales or Amphisphaeriales (Pyrenomycetes, Ascomycota). The classification of the anamorph is based primarily on conidial morphology i.e. the number of cells, and appendage type. However, UV irradiation can convert typical conidia of Pestalotiopsis microspora (5 celled, 2-3 apical and 1 basal appendage) into fungal biotypes that bear a conidial resemblance to the genera Monochaetia and Truncatella. The single cell cultures of putants retain 100% homologies to 5.8S and ITS regions of DNA in the wild type, suggesting that no UV induced mutation occurred in these regions. These results call to question the stability of conidial morphology and taxonomic reliance on this characteristic for this group of fungi. Therefore, a molecular phylogenetic approach was used to clarify their taxonomic relationships. Teleomorphs of these endophytes were previously placed in either Xylariales or Amphisphaeriales. Based on parsimony analysis of partial 18S rDNA sequences for selected anamorphic and teleomorphic taxa in Amphisphaeriaceae, this research supports the placement of these fungal genera in the order Xylariales sharing a common ancestor with some taxa in Xylariaceae. The 18S rDNA region is helpful in resolving phylogenetic relationships at or above the ordinal level for these sampled taxa. Parsimony analyses of the ITS1-5.8S-ITS2 sequences suggest that selected genera in Amphisphaeriaceae are a monophyletic group sharing the synapomorphic characteristic of an anamorphic state which has an annellidic holoblastic type of conidiogenesis in acervuli of textura angularis cells. These taxa were delineated into 5 groups somewhat consistent with previous classical classifications. Many of the taxa in this group of fungi are either closely related or identical according to the 97-100 % homologies of the complete ITS1-5.8S-ITS2 regions. Results reflect a correlation between host and the geographical origin for some clades of the inferred tree. A majority of temperate taxa are with gymnosperm hosts, and one group is on tropical palm. Taxol production was detected only in anamorphs, particularly in Seiridium, and for genera on palm or yew, but this may reflect sampling bias. TAXONOMY, MOLECULAR PHYLOGENY AND TAXOL PRODUCTION IN SELECTED GENERA OF ENDOPHYTIC FUNGI by Jeerapun Worapong A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Plant Pathology MONTANA STATE UNIVERSITY Bozeman, Montana May 2001 Ii v no 5- APPROVAL of a dissertation submitted by Jeerapun Worapong This dissertation has been read by each member of the dissertation committee and has been found to be satisfactory regarding content, English usage, format, citations, bibliographic style, and consistency, and is ready for submission to the College of Graduate Studies. (Date) Approved for the Departmerit of Plant Science and Plant Pathology Dr. Norman F. Weeden / / j (Date) Approved for the College of Graduate Studies Dr. Bruce R. McLeod O/ (Date) ■Ill STATEMENT OF PERMISSION TO USE In presenting this dissertation in partial fulfillment o f the requirements for a doctoral degree at Montana State University, I agree that the Library shall make it available to borrowers under rules of the Library. I further agree that copying o f this dissertation is allowable only for scholarly purposes, consistent with “fair use” as prescribed in the U.S. Copyright Law. Requests for extensive copying or reproduction of this dissertation should be referred to Bell & Howell Information and Learning, 300 North Zeeb Road, Ann Arbor, Michigan 48106, to whom I have granted “the exclusive right to reproduce i and distribute my dissertation in and from microform along with the non-exclusive right to reproduce and distribute my abstract in any format in whole or in part.” ACKNOWLEDGMENTS I would like to express my deepest appreciation to my major advisor, Dr. Gary A. Strobel for kindly providing me with the opportunity to pursue a doctorate degree at the Montana State University-Bozeman, USA as well as all facilities, professional input and his support and guidance regarding creativity and independent thinking. I also gratefully appreciate Dr. Cathy Cripps for all her support, guidance and encouragement in conducting this dissertation and help in becoming a good mycologist and manuscript writer. To Dr. Don E. Matbre, Dr. Mark Young, Dr. Matt Lavin and Dr. Anneke M. Metz, I am greatly indebted to you for serving on my committee and for their advice during my class work as well as for fruitful discussions and suggestions concerning my dissertation projects. I would like to express a special acknowledgement to Dr. Joseph W. Spatafora, Oregon State University, for providing the 18S rDNA sequences of some orders in Pyrenomycetes, Dr W. M. Hess, Brigham Young University, for providing all scanning electron microscopic pictures of atypical conidia of Putants, Dr. Joe Sears, Montana State University for performing the MS test and Ms. Mary Bateson, o f the MSU sequencing lab for providing all sequence data shown in this dissertation. I wish to state my sincerely appreciation to Dr. Eugene J. Ford, Dr. Dave Long for their valued advice for this dissertation research. Special thanks also to my fellow graduate students and all members of the entire Department of Plant Pathology at MSU for their warmest friendship and their help in the past years. I am thankful for the financial support provided by Mahidol University, the Government of Thailand and P.E.O. International Peace Scholarship. Finally I would like to extend my deepest appreciation to my mother, Prapai Worapong, and my elder brother, Worapoj Worapong, for their love and support, and to Dr. Chaufah Thongthai and Dr. Nongluksna Sriubolmas and friends in Thailand for their encouragement. TABLE OF CONTENTS Page LIST OF TABLES................................................................................................................... .. LIST OF FIGURES........................................................................................................................ ABSTRACT.............................................................................................................................. .. 1. LITERATURE REVIEW...................................................................................................... Endophytic Fungi.................................................................................................................. Evolution o f Endophytes from Latent pathogens to Mutualistic Symbiosis.................................................................................................. Strategies of Mutualism in Endophytes.... ...................................................................4 Taxonomy of Endophytes...............................................................................................4 Rainforest Endophytes................................................................. 5 The Fungal System..................................................................................................................9 The Hierarchical Classification..................................................................................... 9 Definition o f Fungi....................................................................................................... Classification of Fungi.................................................................................................. 11 Nomenclature....................................... ......................................................................... 13 A Dual Nomenclature....................................................................................................13 Classical Classification................................................................................................. 14 Advantages of Morphological Characteristic Data....................................................16 Molecular Phylogeny.......................... 16 Fungal molecular Data.................................................................................................. 18 Advantages of Molecular Data in Phylogenetic Studies...........................................20 The concept of Species in Fungi......................................................................................... 21 Morphological Species Concept (MSC)...............................................................;.. .22 Biological Species Concept (BSC)............................... 23 Phylogenetic Species Concept (PSC)......................................................................... 23 Advantages and Disadvantages of MSC, BSC, PSC................................................24 Taxonomy o f Amphisphaeriaceae.....................................................................................
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