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Identification and Pathogenicity of Bursaphelenchus Species (Nematoda: Parasitaphelenchidae)

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Identification and Pathogenicity of Bursaphelenchus Species (Nematoda: Parasitaphelenchidae) Identification and pathogenicity of Bursaphelenchus species (Nematoda: Parasitaphelenchidae) Hongmei Li There is a dim light at the other end of the tunnel The tunnel brightens more as I draw closer Fear, my worst enemy wants me to give up Hope my most valuable companion keeps me moving on Taking one step ahead is not easy but I shall overcome For I believe that all the suffering is not put to waste For there is a dim light at the other end of the tunnel -Nathan Gwira- PROMOTER: PROF. DR IR MAURICE MOENS GHENT UNIVERSITY, FACULTY OF BIOSCIENCE ENGINEERING, DEPARTMENT OF CROP PROTECTION INSTITUTE FOR AGRICULTURAL AND FISHERIES RESEARCH PROF. DR IR LUC TIRRY GHENT UNIVERSITY, FACULTY OF BIOSCIENCE ENGINEERING, DEPARTMENT OF CROP PROTECTION DEAN: PROF. DR IR HERMAN VAN LANGENHOVE RECTOR: PROF. DR PAUL VAN CAUWENBERGE HONGMEI LI IDENTIFICATION AND PATHOGENICITY OF BURSAPHELENCHUS SPECIES (NEMATODA: PARASITAPHELENCHIDAE) THESIS SUBMITTED IN FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR (PHD) IN APPLIED BIOLOGICAL SCIENCES HONGMEI LI IDENTIFICATIE EN PATHOGENICITEIT VAN BURSAPHELENCHUS SPECIES (NEMATODA: PARASITAPHELENCHIDAE) COVER FIGURE: PINE WILT DISEASE IN PURPLE MOUNTAIN, NANJING, CHINA IN 1985. LI, H. (2008). IDENTIFICATION AND PATHOGENICITY OF BURSAPHELENCHUS SPECIES (NEMATODA: PARASITAPHELENCHIDAE). PHD THESIS, GHENT UNIVERSITY, GHENT, BELGIUM. ISBN-NUMBER: 978-90-5989-235-4 THE AUTHOR AND THE PROMOTERS GIVE THE AUTHORIZATION TO CONSULT AND TO COPY PARTS OF THIS WORK FOR PERSONAL USE ONLY. EVERY OTHER USE IS SUBJECT TO THE COPYRIGHT LAWS. PERMISSION TO REPRODUCE ANY MATERIAL CONTAINED IN THIS WORK SHOULD BE OBTAINED FROM THE AUTHOR ACKNOWLEDGEMENTS My thesis work has received the financial support by a PhD scholarship of the Flemish Interuniversity Council-University Development Cooperation (VLIR-UOS) of Belgium. Scientifically, my work has benefited from the Faculty of Bioscience Engineering of Ghent University, the Department of Plant Pathology of Nanjing Agricultural University, China, the Institute for Agricultural and Fisheries Research (ILVO), Belgium and the Scottish Crop Research Institute (SCRI). It is my privilege to express my sincere thanks to all who helped me directly or indirectly during my four-year study. First of all I wish to express my sincere appreciation and gratitude to my supervisor Professor Dr ir. Maurice Moens for his assistance in all steps in the preparation of this thesis. Under his guidance and help, I successfully overcame many difficulties and learnt a lot about how to work with nematodes. I also express my sincerest gratitude to his wife Monique who gave me a lot of care during my stay in Belgium. Their love to me gave me the strength to pick up my confidence. I wish to extend my deepest thanks to my co-promoter Prof. Dr ir. Luc Tirry for accepting me as his student and for providing me with valuable suggestions and help. I am thankful to my local supervisor Prof. Dr Maosong Lin for his support in my field experiments. I thank the members of the reading committee, Profs Roland Perry and John Jones, for the valuable time they devoted to my thesis, and to Prof. Dr ir. Walter Viaene, Prof. Dr ir. Erik Van Bockstaele and Prof. Dr ir. Patrick De Clercq for evaluating my work. I wish to express my gratefulness to Prof. Dr John Jones (SCRI) for providing me a unique opportunity to work under his guidance. I deeply thank him for his encouragement to improve my scientific abilities and his support in writing. I also wish to acknowledge Profs Jones and Moens who, through a common project sponsored by the Royal Society (UK), gave me the chance to visit SCRI during the past four years. I extend special thanks to Dr Mark Philips for helping me to analyse the sequence data and Dr Vivian Block for her valuable suggestions. I appreciate very much the staff of the Nematology research group who gave me a pleasant feeling in the lab. Special thanks go to my Scottish landlords, Ms. Anna Anderson and Mr. John Stoa, who gave me a warm family life when being in Dundee. i I thank the Flemish Interuniversity Council (VLIR) for providing me with a grant to attend the Postgraduate International Nematology Course (PINC) organised at Ghent University, which was a prerequisite for this PhD program. Prof. Moens, the director of PINC, and all PINC professors are acknowledged. Special thanks go to the coordinator of the course Ms. Nic Smol and the secretaries Ms. Kristien Ballegeer and Ms. Inge Dehennin. During the time I worked at ILVO - Unit Plant - Crop Protection (former CLO), I experienced the kindness from all staff. I extend my thanks to the ILVO- Crop Protection staff and colleague students in the department. I am grateful to Lieven Waeyenberge, Nancy de Sutter, Nicole Viaene, Wim Wesemael, Bart Cottyn, Rachid Tahzima, Etienne De Smet, Hubert Van De Sype, Erwin De Poorter, Pieter Callebert, Tine Maes, Steve Baeyen, Steven Cools and John Witters, who helped me in one way or another; I appreciate their friendship during my stay. I would like to give my thanks to Prof. Manuel Mota and Paulo Vieira from the Nematology Lab-ICAM, Universidade de Évora (Portugal) for providing important literature. I also thank Prof. Dr Etienne Geraert from Ghent University for providing extra literature needed for research work. I address special thanks to Dr Taisei Kikuchi from the Forestry and Forest Products Research Institute (Japan) for the good cooperation in the experimental work. I would like to thank Mr. Peiyin Shen from the Inspection and Quarantine for Exit and Entry of Nanjing (China) for collecting nematode samples from wood packaging and preparing wood logs. Dr Fuyuan Xu from the Forestry Academy of Jiangsu Province (China) is acknowledged for providing me with pine trees. I want to give my very special thanks to my classmate Mr. Trinh Quang Phap from Institute of Ecology and Biological Resources of Vietnam. His company in the lab gave me the most valuable help; I appreciate his friendship deeply. I also want to thank my other Vietnamese friends, Ms. Nguyen Thi Bao Anh, Nguyen Thi Ngoc Anh, Duong Thi Minh Nguyet, Nguyen Thi Tuyet and Mr. Lee Hong Phouc for their company in my life in Ghent and the joyful time they gave to enjoy Vietnamese culture. I am indebted to the directors of Nanjing Agricultural University (China) who gave me the opportunity to study abroad. Special thanks go to my colleagues in the Department of Plant Pathology who took care of my lab during my absence; I especially thank my colleagues Prof. Dr Jianhua Xu and Ms. Peng Fu. ii I also thank my students in the lab for their understanding and hard work during my absence in the lab for a long time. I would like to express my deep gratitude to all my Chinese friends both in China and in Ghent, of which I cannot add their entire name here. Finally I want to thank for the love I received from my father whenever and wherever I was. My deepest thanks go to my twin sons Jiakun (Shark) and Jiapeng (Eagle), whose love let me overcome the most difficult time in my life. I am painfully sorry for my absence far from them for a long time whilst I could not see them growing up. Their eternal love to me has encouraged me to finish the study; finally I shall join them being a happy mother. iii TABLE OF CONTENTS Acknowledgements……...…………………………………………………….………………i Table of contents……………………………………………………………………………...iv List of figures …………………………………………………………..……………..….....viii List of tables.……………………………………………………….……………….……....xiii Chapter 1: General Introduction………………………….…………………..……..………1 Chapter 2: Bursaphelenchus species: biology, identification and pathogenicity….………7 2.1 Pine wood nematode Bursaphelenchus xylophilus ……………………………………...8 2.1.1 History and distribution ………………………………………………………….....8 2.1.2 Biology …………………………………………………………………….………10 2.1.2.1 Life cycle and behaviour …………………………………………………..….11 2.1.2.2 Host plants ………………………………………………………………….…13 2.1.2.3 Insect vectors ………………………………………………………………….13 2.1.2.4 Movement and dispersal ………………………………………………………14 2.1.3 Management and control …………………………………………………………..15 2.2 Taxonomy and identification of Bursaphelenchus species …………………………….16 2.2.1 Taxonomical position of genus Bursaphelenchus …………………………………16 2.2.2 The species of genus Bursaphelenchus ……………………………………………16 2.2.3 The taxonomy of the genus Bursaphelenchus ……………………….………………...22 2.2.4 Species identification within Bursaphelenchus ………………………………….........23 2.2.4.1 Morphological methods ………………………………………………………23 2.2.4.2 Protein based methods ………………………………………………………..23 2.2.4.3 DNA based approaches………………………………………………………..24 2.2.4.3.1 DNA-hybridization methods ……………………………………………...24 2.2.4.3.2 PCR-based methods ………………………………………………………24 2.3 Pathogenicity of Bursaphelenchus xylophilus ……...………………………………….26 2.3.1 Development of disease …………………………………………………………...27 2.3.2 Factors important in development of pine wilt disease……………………………28 2.3.2.1 Nematode populations ………………………………………………………...28 2.3.2.2 Host plants …………………………………………………………………….29 2.3.2.3 Environmental factors …………………………………………………………29 2.3.3 Mechanism of pathogenicity ………………………………………………………30 2.3.3.1 Causes of pine wilt disease ……………………………………………………31 2.3.3.1.1 Nematode as the sole pathogen ……………………………………….…..31 2.3.3.1.2 Nematode and bacterial disease complex..………………………………...31 2.3.3.2 Pathogenesis for pine wilt disease …………………………………….………32 2.3.3.2.1 Enzymes.………………………………………………………………..…32 2.3.3.2.2 Chemicals.………………………………………………………………....33 2.3.3.2.3 Phytotoxins…………………………………………………………….….33 2.3.3.2.3.1 Production of phytotoxins
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