Pine Wilt Disease in Portugal
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Bo Guang Zhao • Kazuyoshi Futai • Jack R. Sutherland • Yuko Takeuchi Editors Pine Wilt Disease Bo Guang Zhao • Kazuyoshi Futai • Jack R. Sutherland • Yuko Takeuchi Editors Pine Wilt Disease With 139 Figures, Including 26 in Color Bo Guang Zhao, Ph.D. Professor College of Forest Resources and Environment Nanjing Forestry University 159 Longpan Road Nanjing 2110037, People’s Republic of China Kazuyoshi Futai, Ph.D. Professor Laboratory of Environmental Mycoscience Graduate School of Agriculture, Kyoto University Kitashirakawa Oiwake-cho, Sakyo-ku Kyoto 606-8502, Japan Jack R. Sutherland, Ph.D. Former Principal Research Scientist Pacifi c Forestry Centre 506 West Burnside Road, Victoria, British Columbia V8Z 1M5, Canada Yuko Takeuchi, Ph.D. Assistant Professor Laboratory of Environmental Mycoscience Graduate School of Agriculture, Kyoto University Kitashirakawa Oiwake-cho, Sakyo-ku Kyoto 606-8502, Japan Library of Congress Control Number: 2008926209 ISBN: 978-4-431-75654-5 Springer Tokyo Berlin Heidelberg NewYork e-ISBN: 978-4-431-75655-2 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifi cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfi lms or in other ways, and storage in data banks. The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specifi c statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Springer is a part of Springer Science+Business Media springer.com © Springer 2008 Printed in Japan Typesetting: SNP Best-set Typesetter Ltd., Hong Kong Printing and binding: Kato Bunmeisha, Japan Printed on acid-free paper We dedicate this book to Dr. Y. Tokushige, Dr. T. Kiyohara, Dr. W.R. Nickle, Dr. Y. Mamiya, and all other pioneers in this fi eld. Foreword At the turn of the 20th Century, Japanese foresters began to notice the widespread mortality of pines around the port city of Nagasaki. For several decades the mortal- ity spread northward through the main island of Honshu. At the time was thought that wood boring beetles, prevalent in the dead trees, were the cause of the mortal- ity. The epidemic of pine mortality continues to this day; however, the nature of the investigation into the epidemiology of the disease changed with the determina- tion by Kiyohara and Tokushige in 1971, that the causal agent to the rapid wilting of Japanese red and Japanese black pines was not the beetle but rather a species of nematode vectored by the beetle. This discovery was not anticipated as nematodes were not considered the casual agents of tree disease. Indeed, a quick review of forest pathology texts from that time would provide a paucity of signifi cant information about nematodes. The only nematode-caused tree disease to be found in the literature of the time was red ring disease of coconut palm caused by the weevil-vectored nematode, Bursaphelenchus cocophilus. With the identifi cation of the pine wood nematode, serious research on the biology and ecology of pine wilt began in Japan. A few years later, at the sugges- tion of visiting researchers from Japan, pine wood nematodes were recovered from the wood of a dead Austrian pine in Columbia, Missouri, USA. A fi rst thought to be an introduction of the nematode into North America it was soon realized that the nematode was native to North America and was likely introduced into Japan in the early 1900s. Human activity has resulted in the intentional and accidently distribution of plant and animal species beyond their historical geographical distributions. Most unin- tentional introductions fail to establish in their new environment; however, there have been many striking examples of ecosystems being threaten by the establish- ment of an exotic organism. Among the dramatic examples in forest ecosystems are chestnut blight, Dutch elm disease, gypsy moth and the hemlock woolly adelgid. The introduction of the pine wood nematode into Japan and its subsequent spread to China, Korea and more recently Portugal is yet another example. VI Foreword VII The research community continues to make impressive strides in understanding the complex relationships that govern the interactions between the nematode, the beetle, the tree and associated microorganisms. The system is a beautifully syn- chronized series of interactions that involves numerous cross species signals that act to synchronize tree decline, beetle colonization, nematode development and nematode attraction to newly-formed adult beetles. This volume represents the state of knowledge of the pine wilt system in eastern Asia and the efforts to control its spread. As such it will become a valuable resource for researchers, resource managers and students. Marc J. Linit Professor of Entomology and Associate Dean for Research and Extension University of Missouri, Columbia, Missouri, USA IUFRO Coordinator Working Party 7.02.10—Pine Wilt Disease Preface The year 1905 witnessed the fi rst description of pine wilt disease at Nagasaki on the island of Kyushu in Japan, but the pine wood nematode, Bursaphelenchus xylophilus, was not identifi ed as the causal agent of the disease until 1971, when Kiyohara and Tokushige demonstrated the pathogenicity of the nematode by inocu- lating 25-year-old Pinus densifl ora. The analysis of DNA from several studies indicates that the pine wood nematode was introduced to Japan from the United States. Since the fi rst outbreak on the island of Honshu at Aioi in 1914, the disease has been found in North America (Canada, the United States, and Mexico), in East Asia (Japan, Korea, and China), and in Europe (Portugal). It has now become a worldwide threat to forest ecology and international trade. Thanks to vigorous work and unremitting efforts devoted to research into pine wilt disease by scientists around the world, we now have a better understanding of the disease, e.g., systematics, diagnostics of the nematode, and the biology, ecology and modeling of the insect vectors, physiology and resistance of the host tree, and interactions between the nematode and its associated microbe. However, more mysteries behind the symptoms remain to be explained. For example, many scien- tists agree that toxins are involved in the disease, but the origin or origins of certain toxins are not entirely clear. Reports have shown that axenic B. xylophilus lost its pathogenicity and did not result in wilting symptoms when host seedlings were inoculated with it. Meanwhile, inoculation with a mixture of pathogenic bacteria carried by the nematode and the axenic nematode resulted in wilt and death of seedlings in Pinus thunbergii. Moreover, mutualistic symbiosis between the nema- tode and the bacteria carrying it was discovered. A hypothesis was then proposed, namely, that pine wilt disease was a complex one caused by both the nematode and its associated pathogenic bacteria, which sheds light on a new direction for research. I think better understanding of the etiology of the disease will pave the way to stopping its spread. Therefore, further basic research is needed in interac- tions between B. xylophilus, the bacterial pathogens, and the molecular biology of pathogenicity in the disease. However, some scientists, on the basis of their own experiments, hold opposing opinions, contrary to the above-mentioned hypothesis. I think that is quite normal. Debate among those with differing points of view will VIII Preface IX accelerate the development of a better understanding of certain scientifi c issues. We welcome critical reviews from readers of all the contents of this book. When Dr. K. Futai paid a visit to China in 2006, we discussed the possibility of compiling a book on pine wilt disease for the purpose of providing useful assistance for researchers, university graduates, and foresters, and both of us believed that it was the right time to compile a book integrating recent research progress and criti- cal assessment of the state of our understanding of the disease. Since then, many prestigious scientists knowledgeable about the disease and the research have been invited to join our efforts by contributing their intelligence and wisdom to the compilation of the book. I would like to thank all the contributors to the book whether or not they are listed as authors, and I appreciate those who conducted most of the fi eld work and scientifi c instrument analyses. Special acknowledgments are also given to the spon- sors and providers of funds for their support for the research resulting in publication of this book. My project team thanks the Natural Science Foundation of China (Key Projects Nos. 30430580 and 30030110), the State Forestry Administration, the People’s Republic of China (Project No. 20070430), and the Nanjing Forestry University for their fi nancial support of our research. Nanjing, May 2008 Professor Bo Guang Zhao Nanjing Forestry University People’s Republic of China Contents Foreword . VI Preface . VIII Contributors . XIII Color Plates . XVII Part I Historical Overview. 1 1 Introduction . 2 2 Pine Wilt in Japan: From First Incidence to the Present . 5 Kazuyoshi Futai 3 A Brief Overview of the Pine Wood Nematode and Pine Wilt Disease in Canada and the United States . 13 Jack R. Sutherland 4 Pine Wilt Disease in China . 18 Bo Guang Zhao 5 Pine Wilt Disease in Korea . 26 Sang-Chul Shin 6 Pine Wilt Disease in Portugal . 33 Manuel M. Mota and Paulo C. Vieira 7 Concluding Remarks . 39 Part II Pine Wood Nematode . 41 8 Introduction . 42 9 Taxonomy and Systematics of the Nematode Genus Bursaphelenchus (Nematoda: Parasitaphelenchidae) . 44 Natsumi Kanzaki 10 Parasitism Genes of the Pine Wood Nematode . 67 Taisei Kikuchi X Contents XI 11 Embryology and Cytology of Bursaphelenchus xylophilus . 81 Koichi Hasegawa and Johji Miwa 12 Population Ecology of Bursaphelenchus xylophilus . 105 Shuhei Takemoto 13 Transmission Biology of Bursaphelenchus xylophilus in Relation to Its Insect Vector .