Advances in Genetics, Genomics and Controlof Rice Blast Disease

Guo-Liang Wang · Barbara ValentEditors

Advances in Genetics,Genomics and Controlof Rice Blast Disease

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Editors

Prof. Guo-Liang WangOhio State UniversityDepartment of Plant Pathology2021 Coffey RoadOhio OH 43210-1087USAwang.620@osu.edu

Prof. Barbara ValentKansas State UniversityDepartment of Plant Pathology4108 ThrockmortonManhattan KS 66506-5502USAbvalent@ksu.edu

ISBN 978-1-4020-9499-6 e-ISBN 978-1-4020-9500-9

DOI 10.1007/978-1-4020-9500-9

Library of Congress Control Number: 2008940153

c© Springer Science+Business Media B.V. 2009No part of this work may be reproduced, stored in a retrieval system, or transmittedin any form or by any means, electronic, mechanical, photocopying, microfilming, recordingor otherwise, without written permission from the Publisher, with the exceptionof any material supplied specifically for the purpose of being enteredand executed on a computer system, for exclusive use by the purchaser of the work.

Front Cover: Neck blast. Photo by Barbara Valent from the field trip at the 3rd International Rice BlastConference in Japan, 2003.Back Cover: Projected confocal image of Magnaporthe oryzae invading rice sheath epidermal cells.The fungus expresses cytoplasmic EYFP (enhanced yellow fluorescent protein; pseudocolored blue) andnuclear-localizing mRFP (monomeric red fluorescent protein) at 36 hour post inoculation. The embosseffect was added using PowerPoint. Image from Chang Hyun Khang and Barbara Valent.

Printed on acid-free paper

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Preface

Rice blast, caused by the fungus Magnaporthe oryzae, is one of the first recordeddiseases of rice. It was known as rice fever disease in China as early as 1637 and wasreported as Imochi-byo in Japan in 1704. As the rice production expanded throughAsia, Latin America, and Africa over the last few centuries, the disease followedand is now found in over 85 countries worldwide. M. oryzae attacks all parts of therice plant causing losses upwards of hundreds of millions of tons of rice grain annu-ally. Such losses have lead to rice shortages in many developing countries in recentyears, making effective control of this devastating disease imperative for global foodsecurity and social well-being.

Fortunately, the genomes of both rice and M. oryzae were completely sequencedwithin the last five years. Due to unrestrained public availability of the genomesequences, genetic stocks, and improved research tools, this pathosystem has becomethe premier model for understanding the molecular basis of plant-fungal interac-tions. Since the 3rd International Rice Blast Conference (IRBC) held on September11–14, 2002 in Tsukuba, Japan, significant advancements have emerged in ourunderstanding of this disease and technologies available for analysis. To reviewthe recent progress and discuss the future of rice blast research, the 4th IRBC wasconvened on October 10–14, 2007 in Changsha, the capital of Hunan province,China.

The premise for having the conference in China is threefold. First, China is thelargest rice producing country in the world with rice being the nutritional staplefor over 60% of its 1.3 billion people. Sustainable rice production is essential forthe economic and social stability of this country. Second, rice blast continues tocause significant yield losses in China, with many newly-bred rice varieties los-ing their genetic resistance within years of being released. Yield losses in a fewelite hybrid rice lines were reported to be as high as 80–90% in recent years inhard hit regions. Finally, Chinese scientists began actively working on rice blastabout 50 years ago. In the time since, they have made considerable achievementsin selecting blast resistant germplasm for breeding programs, monitoring rice blastpopulations and epidemics, and developing effective cultural and chemical controlprotocols. Their recent progress in the molecular analysis of the defense response inrice and conversely pathogenicity in M. oryzae is laudable. China now has arguablythe largest cohort of scientists in the world focusing on this disease. Therefore, the

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proposal to hold the 4th IBRC in China was well received and fully supported bythe Rice Blast Executive Committee.

The conference was successfully held in the new, well-appointed Vaya Interna-tional Hotel. More than 250 attendees from 20 countries convened, making it thelargest IRBC to date. There were 78 oral presentations and 64 poster presentations.Of special note, many postdoctoral fellows and graduate students, representing thefuture leaders of this field, attended the conference. A unique and memorable eventwas the award ceremony at the closing banquet in West Lake Restaurant. Distin-guished Research Awards were presented to ten scientists who devoted their entirecareer to rice blast research. Eight of these distinguished scientists were able toattend the conference and accepted the recognition plaque prepared by the RiceBlast Executive Committee and the 4th IRBC Organizing Committee. The confer-ence concluded with dramatic traditional Chinese dances and performances. By allaccounts, the 4th IRBC was a success and once again united the international com-munity of scientists studying this ancient and devastating disease of rice. We nowlook forward to hearing the next major advances and insights shared when rice blastresearch community assembles in Arkansas, USA in 2010.

Columbus, OH, USA Guo-Liang Wang

Acknowledgements

We would like to thank the following institutions and companies for their financialsupport that made the 4th IRBC possible:

Hunan Agricultural UniversityChina Agricultural UniversityYunnan Agricultural University,Chinese Society for Plant PathologyHunan Society for Plant PathologyPre-National Key Laboratory of Plant Germplasm Generation and UtilizationOhio State UniversityMinistry of Science and Technology of the People’s Republic of China (973Project)National Natural Science Foundation of ChinaMinistry of Agriculture of the People’s Republic of China (948 project)Department of Science and Technology of Hunan ProvinceSyngenta Agroservices Asia Ltd.Bayer CropScience Inc.Beckman Coulter, Inc.

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Contents

Current Status and Future Prospects for Research on Blast Resistancein Rice (Oryza sativa L.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1Gurdev S. Khush and K.K. Jena

Part I Pathogen Genetics, Genomics and Molecular Biology

The PMK1 MAP Kinase Pathway and Infection-Related Morphogenesis . . 13Shengli Ding, Xiaoying Zhou, Xinhua Zhao and Jin-Rong Xu

Surface Sensing and Signaling During Initiation of Rice-Blast Disease . . . . 23Hao Liu, Ravikrishna Ramanujam and Naweed I. Naqvi

Studies on Autophagy Machinery in Magnaporthe oryzae . . . . . . . . . . . . . . . 33Fu-Cheng Lin, Xiao-Hong Liu, Jian-Ping Lu and Tong-Bao Liu

Secretion Property and Gene Expression Pattern of a Putative FeruloylEsterase in Magnaporthe grisea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41Xiang-zi Zheng, Jie Zhou, Chen-zeng Lin, Xiong-jie Lin, Lan Lan,Zong-hua Wang and Guo-dong Lu

Functional Analysis of Two Laccase Genes in Magnaporthe grisea . . . . . . . . 51Xin Chen, Wende Liu, Chuanzhi Zhao, Shuji Liu, Minoo Razee,Guo-dong Lu and Zonghua Wang

The Significance of Nitrogen Regulation, Source and Availabilityon the Interaction Between Rice and Rice Blast . . . . . . . . . . . . . . . . . . . . . . . . 59Nicole M. Donofrio, Thomas K. Mitchell and Ralph A. Dean

Protein Chips and Chromatin Immunoprecipitation – EmergingTechnologies to Study Macromolecule Interactions in M. grisea . . . . . . . . . . 73Thomas K. Mitchell, Ralph A. Dean, Jin-Rong Xu, Heng Zhu, Yeon Yee Ohand Hee-Sool Rho

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Cellular and Molecular Analyses of Biotrophic Invasion in Rice BlastDisease . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83Prasanna Kankanala, Gloria Mosquera, Chang Hyun Khang,Guadalupe Valdovinos-Ponce and Barbara Valent

Isolation and Functional Analysis of Putative Effectors fromMagnaporthe oryzae Using Integrated Genomic Approaches . . . . . . . . . . . . . 93Songbiao Chen, Malali Gowda, R.C. Venu, Pattavipha Songkumarn,Chan Ho Park, Maria Bellizzi, Daniel J. Ebbole and Guo-Liang Wang

Searching for Effectors of Magnaporthe oryzae: A Multi-FacetedGenomics Approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105Ryohei Terauchi, Joe Win, Sophien Kamoun, Hideo Matsumura,Hiromasa Saitoh, Hiroyuki Kanzaki, Kentaro Yoshida, Matt Shenton,Thomas Berberich, Shizuko Fujisawa, Akiko Ito, Yoshitaka Takanoand Yukio Tosa

Developing Resources for Analysis of Secreted Proteinsfrom Magnaporthe oryzae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113Guo-dong Lu, Hanno Wolf, Yue Shang, Cristina Filippi, Kiran Bhattarai,Dan Li and Daniel J. Ebbole

Part II Host Defense Genetics, Genomics, Molecular Biology

Functional and Evolutionary Analysis of the Pi2/9 Locus in Rice . . . . . . . . . 127Bo Zhou and Guo-Liang Wang

Understanding the Co-evolution of the Rice Blast Resistance GenePI-TA and Magnaporthe oryzae Avirulence Gene AVR-PITA . . . . . . . . . . . . . 137Yulin Jia, Xueyan Wang, Stefano Costanzo and Seonghee Lee

Genetic and Molecular Analyses of Blast Resistance in a Universal BlastResistant Variety, Digu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149Xuewei Chen, Junjun Shang, Cailin Lei, Jichen Xu, Shigui Liand Lihuang Zhu

Map-Based Cloning and Breeding Application of a Broad-SpectrumResistance Gene Pigm to Rice Blast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161Yiwen Deng, Xudong Zhu, Jing Xu, Hongqi Chen and Zuhua He

Rac GTPase and the Regulation of NADPH Oxidase in Rice InnateImmunity Response . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173Hann Ling Wong, Tsutomu Kawasaki and Ko Shimamoto

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Rice Guanine Nucleotide Exchange Factors for Small GTPase OsRac1Involved in Innate Immunity of Rice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179Tsutomu Kawasaki, Keiko Imai, Hann Ling Wong, Yoji Kawano,Keita Nishide, Jun Okuda and Ko Shimamoto

Role of Ethylene, Abscisic Acid and MAP Kinase Pathwaysin Rice Blast Resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185Tameka A. Bailey, Xiangjun Zhou, Jianping Chen and Yinong Yang

Bacterial Determinants and Host Defense Responses UnderpinningRhizobacteria-Mediated Systemic Resistance in Rice . . . . . . . . . . . . . . . . . . . 191David De Vleesschauwer and Monica Hofte

Part III Pathogen Population and Disease Control

World Population Structure and Migration of the Rice Blast Fungus,Magnaporthe oryzae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209Didier Tharreau, Isabelle Fudal, Dodelys Andriantsimialona,Santoso, Dwinita Utami, Elisabeth Fournier, Marc-Henri Lebrunand Jean-Loup Notteghem

Examination of the Rice Blast Pathogen Population Diversityin Arkansas, USA – Stable or Unstable? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217J.C. Correll, E.J. Boza, E. Seyran, R.D. Cartwright, Yulin Jia and F.N. Lee

Studies on the Complementary Differential Varieties and LocalPhysiologic Races of Magnaporthe grisea in Sichuan Province . . . . . . . . . . . 229Hongli Ji, Li Shen, Yunjia Xiang, Zhenyu Zhang, Huaming Liao,Linming Luo, Zhen Qin and Yunliang Peng

Blast Race Monitoring for Stable Use of Blast Resistance in Rice . . . . . . . . . 239Shinzo Koizumi

Pathogenicity-Related Compounds Produced by Blast Fungus . . . . . . . . . . . 247Tetsu Tsurushima, Hitoshi Nakayashiki, Yukio Tosa and Shigeyuki Mayama

An Unprecedented Outbreak of Rice Blast on a Newly Released CultivarBRS Colosso in Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 257Anne S. Prabhu, M. Cristina Filippi, Gisele B. Silva, Valacia L. Silva Loboand Orlando P. Morais

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Effect of Magnaporthe grisea on Seed Germination, Yield and Quality ofWheat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267A.S. Urashima, C.R.F. Grosso, A. Stabili, E.G. Freitas, C.P. Silva,D.C.S. Netto, I. Franco and J.H. Merola Bottan

Part IV Identification and Use of Resistance Genes and Breeding Strategies

Field Resistance Expressed when the Pi-ta Gene is Compromisedby Magnaporthe oryzae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281Fleet N. Lee, R.D. Cartwright, Yulin Jia and J.C. Correll

Resistance Genes and Their Effects to Blast in Korean Rice Varieties(Oryza sativa L.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291Young-Chan Cho, Jung-Pil Suh, Ji-Ung Jeung, Jae-Hwan Roh,Chang-In Yang, Myung-Kyu Oh, Jong-Seong Jeon, Im-Soo Choi,Hung-Goo Hwang, Sae-June Yang and Yeon-Gyu Kim

Genetic Analysis of Resistance Against Bacterial Leaf Blight and LeafBlast Disease in the Japanese Rice Cultivar Asominori . . . . . . . . . . . . . . . . . . 305Takashi Endo, Toshiki Nakamura, Junichi Yonemaru, Goro Ishikawa,Masayuki Yamaguchi, Tomomori Kataoka, Koji Nakagomiand Narifumi Yokogami

DNA Marker Analysis of Blast Resistance Genes Pib and Pitain IRRI-Bred Rice Varieties Comparing with Gene Estimationby a Differential System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 315Daisuke Fujita, Leodegario A. Ebron, Nobuya Kobayashiand Yoshimichi Fukuta

Genetic Characterization of Universal Differential Variety SetsDeveloped Under the IRRI-Japan Collaborative Research Project . . . . . . . 325Yoshimichi Fukuta, Donghe Xu, Mary Jeanie T. Yanoria, Aris Hairmansis,Nagao Hayashi and Nobuya Kobayashi

Sequential Planting as a Method of Screening of Durable Resistanceto Rice Blast in Korea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337Jae-Hwan Roh, Byung-Ryun Kim, Se-Weon Lee, Young-Chan Cho,Dong-Soo Ra, In-Seok Oh and Seong-Sook Han

Development of a Package of 20 Varieties for Blast Managementon Upland Rice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347Suwarno, Erwina Lubis, Aris Hairmansis and Santoso

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Rice Blast in India and Strategies to Develop Durably Resistant Cultivars 359Mukund Variar, C.M. Vera Cruz, M.G. Carrillo, J.C. Bhatt and R.B.S. Sangar

Breeding Rice Cultivars with Durable Blast Resistance in Colombia . . . . . . 375Fernando Correa Victoria and Cesar Martinez

What it Takes to Achieve Durable Resistance to Rice Blast? . . . . . . . . . . . . . 385Bin Liu, Xiao Yuan Zhu, Shaohong Zhang, Jianli Wu, Seong-Sook Han,Young-Chan Cho, Jae-Hwan Roh, Jan Leach, Yan Liu, Suzette Madamba,Alice Bordeos, Marietta Baraoidan, Isabelita Ona, C.M. Vera Cruzand Hei Leung

Part V Bioinformatics and Database

Magnaporthe grisea Oryza sativa (MGOS) Interaction Database:Community Annotation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 405Kevin A. Greer and Carol Soderlund

ARCHIPELAGO: Towards Bridging the Gap Between Molecularand Genetic Information in Rice Blast Disease Resistance . . . . . . . . . . . . . . . 417E. Ballini, E. Vergne, Didier Tharreau, Jean-Loup Notteghem and J.B. Morel

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427

Awardees of Distinguished Research Awardon Rice Blast at the 4th IRBC

Professor FLEET LEE

Professor Lee, a faculty member at the University of Arkansas, first began work-ing on blast in 1979, and has since received many awards and recognitions. Heestablished the varietal blast screening system that allowed the Arkansas CultivarDevelopment Team to release many successful rice cultivars to farmers. Dr. Leeis described by team members as a “pathologist’s pathologist”, who makes carefulfield observations and then translates them into valuable information for farmersand agricultural scientists. He has been dedicated to understanding field resistanceto blast, and through this dedication, he has optimized flood irrigation practices tohelp farmers to succeed in producing rice even when resistance genes have failed.

Professor LI, DEBAO

Professor Li is one of the pioneer molecular plant pathologists in China. He servedas a deputy governor for Zhejiang province from 1983 to 1993. Professor Li helpedto establish the Institute of Biotechnology at Zhejiang University, and he fosteredand promoted rice blast research there. His latest contribution is to organize theChinese rice blast researchers and initiate the biannual national rice blast conferencein China. Thus, Professor Li has played a major role in promoting rice blast researchthroughout China.

Professor LING, ZHONGZHUAN

Professor Ling has worked with rice blast for over 45 years at the Institute of CropSciences, in the Chinese Academy of Agricultural Sciences. He has published over80 research articles and four books. The neo-isogenic differential lines developedby Professor Ling have been widely used in China and in other countries. Duringhis productive career, Professor Ling received 8 national and institutional awardsfor his contribution to rice blast research. Many highly resistant germplasms wereidentified by Professor Ling and used for breeding cultivars popular in northern

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xvi Awardees of Distinguished Research Award on Rice Blast at 4th IRBC

China. He also made contributions in establishing close collaboration of rice blastresearch between Chinese and Japanese scientists.

Professor LUO, KUAN

Professor Luo is a local hero from Changsha. He studied rice blast for 50 years at theHunan Agriculture University. During this time, he published more than 70 researcharticles on physiological races and virulence of rice blast isolates from Hunan andnearby provinces, and on disease resistance and control. He is the recipient of a firstclass Technology Improvement Award from the Chinese Ministry of Agriculture andfour Science and Technology Progress Awards from Hunan provincial government.

Dr. HAJIME KATO

Dr. Hajime Kato began working on rice blast in 1958, and worked for more than40 years before retiring. Since retirement, he continues his involvement with riceblast. During his distinguished career, Dr. Kato focused on the epidemiology of blastdisease on rice and millets, and on the inoculum source problem for panicle blast.Dr. Kato was one of the pioneers in characterizing the sexual state of the fungus, andhe has been dedicated to understanding the relationship among blast isolates fromdifferent plant species. Dr. Kato’s discovery that the fungus produces microconidiawith some unknown role showed blast researchers how much there still is to learnabout this important disease.

Dr. JEAN-LOUP NOTTEGHEM

One of Dr. Notteghem’s major contributions is the building and maintenance of oneof the best historical, world-wide collections of field isolates of the rice blast fungus.In this process, Dr. Notteghem discovered the most famous fungal strain, GUY11.Dr. Notteghem’s main research theme has been searching for and characterizingupland rice cultivars with high levels of partial resistance and naturally pyramidedresistance genes. This resulted in availability of durably-resistant varieties to breed-ers, such as Moroberekan and IRAT13. Dr. Notteghem was a pioneer in mixingclassical plant pathology and genetics of host-fungus interaction, and in adding inmolecular biology. He now leads a team that uses all the modern genomic toolsavailable to study rice resistance and the rice blast pathogen.

Dr. ANNE SITARAMA PRABHU

Dr. Prabhu began working on blast in Brazil in 1973. He was a pioneer in estab-lishing the first coordinated rice blast program in this large and diverse country. Heestablished a national rice blast nursery, which resulted in the first blast resistantvariety in the field in 1986. Dr. Prabhu received many awards and recognitions,

Awardees of Distinguished Research Award on Rice Blast at 4th IRBC xvii

including the prestigious highest award given to a Brazilian scientist working inthe national research system. He has collaborated with scientists internationally.However, the unique upland rice that is best adapted in Brazil had to be developedin Brazil. Dr. Prabhu’s research resulted in valuable quasi-isogenic lines containing13 resistance genes. Since having resistant rice is not enough for farmers, anothermajor contribution of his was to develop disease management strategies for farmers“to learn to live with rice blast disease”.

Professor PENG, SHAOQUI

Professor Peng graduated from Yangzhou Agriculture University and retired fromthe Institute of Plant Protection, Hunan Academy of Agricultural Sciences fewyears ago. Dr. Peng has studied plant pathology, including rice blast, for 50 years.He has undertaken three projects in the Natural (National) Science Foundation ofChina on “Durable Resistance to Rice Blast”. He has taken part in more than 20national or provincial programs. He has achieved impressive results applying geo-phytopathology to his durable resistance studies on rice blast and plant protection ingeneral. He has been especially active in proposing and promoting study of durableblast resistance at all times within China.

Dr. HIROSHI YAEGASHI

Dr. Yaegashi graduated from Chiba University in Japan in 1965, and he worked onrice blast until his retirement two years ago. Dr. Yaegashi was one of the pioneers incharacterizing the sexual state of the rice blast fungus. Many blast researchers haveoften referenced his famous publication, Yaegashi and Udagawa, 1978, on “Thetaxonomical identity of the perfect state of Pyricularia grisea and its allies.” Dr.Yaegashi published a paper in 1981 that showed that a finger millet pathogen con-tains avirulence genes that correspond to resistance genes in rice. Thus, Dr. Yaegashiled the way in studying the genetics of host specificity in the pathogen.

Professor SHEN, YING

Professor Shen’s career with rice blast spanned over four decades in two provinces,Sichuan and Zhejiang. She contributed significantly to understanding genetic diver-sity, population structure, and geographic distribution of Magnaporthe oryzae inChina and other Asian countries. Professor Shen also was involved in evaluationand utilization of rice germplasm resistant to blast and other major rice diseases.She has authored over 110 papers and received numerous awards from the centraland provincial governments. Professor Shen has been actively involved in manyinternational collaborative projects with scientists in US, Japan, IRRI and Europe.

Contributors

Dodelys Andriantsimialona FOFIFA, URP SCRID, BP230, Antsirabe,Madagascar

Marietta Baraoidan Plant Breeding, Genetics and Biotechnology Division, Inter-national Rice Research Institute, Manila, Philippines

Maria Bellizzi Department of Plant Pathology, The Ohio State University, USA

Thomas Berberich Iwate Biotechnology Research Center, Kitakami, Japan

J.C. Bhatt VPKAS, Almora, Uttarakhand, India

Kiran Bhattarai Department of Plant Pathology & Microbiology, Texas A & MUniversity, College Station, USA 77843

Alice Bordeos Plant Breeding, Genetics and Biotechnology Division, InternationalRice Research Institute, Manila, Philippines

J.H. Merola Bottan Centro de Ciencias Agrarias, Universidade Federal de SaoCarlos, FAPESP (The State of Sao Paulo Research Foundation) Scholars, P.O. Box153, CEP 13600-000 Araras, Sao Paulo, Brazil

E.J. Boza University of Arkansas, Division of Agriculture, Fayetteville, Arkansas72701, USA

M.G. Carrillo International Rice Research Institute, Los Banos, Philippines

R.D. Cartwright Division of Agriculture, Department of Plant Pathology,University of Arkansas, Fayetteville, Arkansas 72701, USA

Hongqi Chen State Key Laboratory of Rice Biology, China National Rice ResearchInstitute, Hangzhou, 310006, China

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xx Contributors

Jianping Chen Department of Plant Pathology and Huck Institutes of Life Sci-ences, Pennsylvania State University, University Park, Pennsylvania 16802, USA

Songbiao Chen Department of Plant Pathology, The Ohio State University, USA

Xin Chen The School of Life Sciences, Fujian Agriculture and ForestryUniversity, Fuzhou 350002, P.R. China

Xuewei Chen State Key Laboratory of Plant Genomics & National Plant GeneResearch Centre (Beijing), Institute of Genetics and Developmental Biology,Chinese Academy of Sciences, Beijing 100101, China; Current address: Departmentof Plant Pathology, University of California, Davis, CA 95616, USA

Young-Chan Cho National Institute of Crop Science, RDA, Suwon 441-857,Korea, yccho@rda.go.kr

Im-Soo Choi National Institute of Crop Science, RDA, Suwon 441-857, Korea

J.C. Correll Division of Agriculture, Department of Plant Pathology, Universityof Arkansas, Fayetteville, Arkansas 72701, USA, jcorrell@uark.edu

Stefano Costanzo USDA-ARS Dale Bumpers National Rice Research Center,Stuttgart, AR72160, USA

C.M. Vera Cruz Plant Breeding, Genetics and Biotechnology Division, Interna-tional Rice Research Institute, Los Banos, Manila, Philippines

David De Vleesschauwer Laboratory of Phytopathology, Faculty of BioscienceEngineering, Ghent University, Coupure links, 653, B-9000 Gent, Belgium

Ralph A. Dean CIFR, Department of Plant Pathology, North Carolina State Uni-versity, Raleigh, NC, 27606

Yiwen Deng National Key Laboratory of Plant Molecular Genetics, Institute ofPlant Physiology and Ecology, Shanghai Institutes for Biological Sciences, Grad-uate School of Chinese Academy of Sciences, Chinese Academy of Sciences,Shanghai 200032, China

Shengli Ding Department of Botany and Plant Pathology, Purdue University, WestLafayette, IN 47907

Nicole M. Donofrio Plant and Soil Science Department, 152 Townsend Hall,University of Delaware, Newark DE, 19716, ndonof@udel.edu

Contributors xxi

Daniel J. Ebbole Department of Plant Pathology and Microbiology, Texas A & MUniversity, College Station, USA 77843, d-ebbole@tamu.edu

Leodegario A. Ebron International Rice Research Institute (IRRI), Los Banos,Laguna, Philippines

Takashi Endo Miyagi Prefectural Furukawa Agricultural Experiment Station(MFAES),Osaki, Miyagi, Japane-mail: tendo@faes.pref.miyagi.jp; endo-ta480@pref.miyagi.jp

Cristina Filippi Department of Plant Pathology & Microbiology, Texas A & MUniversity, College Station, USA 77843

M. Cristina Filippi Embrapa Rice and Beans, Caixa Postal 179, CEP 75375-000,Santo Antonio de Goias, GO, Brazil

Elisabeth Fournier UMR BGPI, CIRAD-INRA-SupAgro.M, Montpellier, France

I. Franco Centro de Ciencias Agrarias, Universidade Federal de Sao Carlos,FAPESP (The State of Sao Paulo Research Foundation) Scholars, P.O.Box 153, CEP13600-000 Araras, Sao Paulo, Brazil

E.G. Freitas Centro de Ciencias Agrarias, Universidade Federal de Sao Carlos.P.O.Box 153, CEP 13600-000 Araras, Sao Paulo, Brazil

Isabelle Fudal UMR BIOGER, INA.PG-INRA, Versailles, France

Shizuko Fujisawa Iwate Biotechnology Research Center, Kitakami, Japan

Daisuke Fujita International Rice Research Institute (IRRI), Los Banos, Laguna,Philippines

Yoshimichi Fukuta Japan International Research Center for Agricultural Sciences(JIRCAS), 1-1, Ohwashi, Tsukuba, Ibaraki 305-8686, Japan

Malali Gowda Department of Plant Pathology, The Ohio State University, USA

Kevin A. Greer Arizona Genomics Computational Laboratory, BIO5 Institute,University of Arizona, Tucson, AZ 85721, USA

C.R.F. Grosso Faculdade de Engenharia de Alimentos, Universidade de Campinas,Cidade Universitaria Zeferino Vaz, CEP 13083-970 Campinas, Sao Paulo, Brazil

Aris Hairmansis Indonesian Center for Rice Research (ICRR), JL. Raya MuaraNo. 25A Ciapus Bogor, Jalan Raya Ciapus 25C, Muara, Bogor, West Java, Indonesia

xxii Contributors

Seong-Sook Han 1Plant Pathology Division, Rural Development Administration,National Institute of Agricultural Science and Technology, RDA, Suwon 441–707,Korea, sshan@rda.go.kr

Nagao Hayashi National Institute of Agrobiological Sciences (NIAS). 2-1-2, Kan-nondai, Tsukuba, Ibaraki 305-8602, Japan

Zuhua He National Key Laboratory of Plant Molecular Genetics, Institute of PlantPhysiology and Ecology, Shanghai Institutes for Biological Sciences, GraduateSchool of Chinese Academy of Sciences, Chinese Academy of Sciences, Shanghai200032, China, zhe@sippe.ac.cn

Monica Hofte Laboratory of Phytopathology, Faculty of Bioscience Engineering,Ghent University, Coupure links, 653, B-9000 Gent, Belgium,Monica.Hofte@UGent.be

Hongli Ji Institute of Plant Protection, Sichuan Academy of Agricultural Sciences,Jingjusilu Road 20, Chengdu 610066, China

Huaming Liao Plant Protection Station under Sichuan Provincial Ministry ofAgriculture, Wuhouci Street 4, Chengdu 610041, China

Hung-Goo Hwang National Institute of Crop Science, RDA, Suwon 441-857,Korea

Keiko Imai Laboratory of Plant Molecular Genetics, Nara Institute of Science andTechnology, Ikoma, 630-0192 Japan

Goro Ishikawa National Agricultural Research Center for TohokuRegion(NARCT), Morioka, Iwate Japan

Akiko Ito Iwate Biotechnology Research Center, Kitakami, Japan

K.K. Jena Senior Scientist, Plant Breeding, Genetics and Biotechnology Division,International Rice Research Institute, C/o IRRI-Korea Office, National Instituteof Crop Science, RDA, 209 Seodun Dong, Suwon 441-857, Republic of Korea,k.jena@cgiar.org

Jong-Seong Jeon Graduate School of Biotechnology and Plant MetabolismResearch Center, Kyung-Hee University, Yongin 446-701, Korea

Ji-Ung Jeung National Institute of Crop Science, RDA, Suwon 441-857, Korea

Contributors xxiii

Yulin Jia USDA-ARS Dale Bumpers National Rice Research Center, Stuttgart,AR 72160, USA, yulin.jia@ars.usda.gov

Sophien Kamoun Sainsbury Laboratory, Norwich, UK

Prasanna Kankanala Department of Plant Pathology, Kansas State University,Manhattan, Kansas, 66506-5502, USA

Hiroyuki Kanzaki Iwate Biotechnology Research Center, Kitakami, Japan

Tomomori Kataoka National Agricultural Research Center for TohokuRegion(NARCT), Morioka, Iwate Japan

Yoji Kawano Laboratory of Plant Molecular Genetics, Nara Institute of Scienceand Technology, Ikoma, 630-0192 Japan

Tsutomu Kawasaki Laboratory of Plant Molecular Genetics, Nara Institute ofScience and Technology (NAIST), 8916-5 Takayama, Ikoma, 630-0192 Nara, Japan,kawasaki@bs.naist.jp

Chang Hyun Khang Department of Plant Pathology, Kansas State University,Manhattan, Kansas, 66506-5502, USA

Gurdev S. Khush Adjunct Professor, University of California Davis, CA 95616,USA, gurdev@khush.org

Yeon-Gyu Kim National Institute of Crop Science, RDA, Suwon 441-857, Korea

Byung-Ryun Kim Chungcheong Nam-Do Agricultural Research and ExtensionServices, Korea

Nobuya Kobayashi International Rice Research Institute (IRRI). DAPO Box7777, Los Banos, Laguna, Metro Manila, Philippines, n.kobayashi@cgiar.org

Shinzo Koizumi National Agricultural Research Center for Tohoku Region,Shimo-Kuriyagawa, Morioka, Iwate, 020-0198 Japan, skoizumi@affrc.go.jp

Lan Lan The School of Life Sciences, Fujian Agriculture and Forestry University,Fuzhou 350002, P.R. China

Jan Leach Colorado State University, Fort Collins, Colorado, USAMarc-Henri Lebrun UMR5240, CNRS-UCB-INSA-Bayer CropScience, Lyon,France

xxiv Contributors

Seonghee Lee USDA-ARS Dale Bumpers National Rice Research Center, Stuttgart,AR72160, USA

F.N. Lee University of Arkansas, Division of Agriculture, Fayetteville, Arkansas72701, USA

Fleet N. Lee University of Arkansas, Rice Research and Extension Center, Stuttgart,AR 72160, USA, fnlee@uark.edu

Se-Weon Lee Plant Pathology Division, National Institute of Agricultural Scienceand Technology, RDA, Suwon, Korea

Cailin Lei Crop Research Institute, Chinese Agricultural Academy of Sciences,Beijing 100085, China

Hei Leung Plant Breeding, Genetics and Biotechnology Division, InternationalRice Research Institute, Manila, Philippines, h.leung@cgiar.org

Dan Li Department of Plant Pathology & Microbiology, Texas A & M University,College Station, USA 77843

Shigui Li Rice Research Institute, Sichuan Agricultural University, Chengdu611130, China

Li Shen Plant Protection Station under Sichuan Provincial Ministry of Agriculture,Wuhouci Street 4, Chengdu 610041, China

Fu-Cheng Lin Biotechnology Institute, Zhejiang University, Kaixuan Road 268,Hangzhou 310029, China, fuchenglin@zju.edu.cn

Chen-zeng Lin The School of Life Sciences, Fujian Agriculture and Forestry Uni-versity, Fuzhou 350002, P.R. China

Xiong-jie Lin The Key Laboratory of Biopesticide and Chemistry Biology, Min-istry of Education, Fujian Agriculture and Forestry University, Fuzhou 350002,P.R. China

Linming Luo Plant Protection Station under Sichuan Provincial Ministry of Agri-culture, Wuhouci Street 4, Chengdu 610041, China

Bin Liu Guangdong Academy of Agricultural Sciences, Guangdong, China

Contributors xxv

Hao Liu Fungal Patho-Biology Group, Temasek Life Sciences Laboratory, 1Research Link, National University of Singapore, Singapore 117604; Institute ofMolecular and Cell Biology, 61 Biopolis Drive, Proteos, Singapore 138673

Shuji Liu The School of Life Sciences, Fujian Agriculture and Forestry University,Fuzhou 350002, P.R. China

Tong-Bao Liu Biotechnology Institute, Zhejiang University, Kaixuan Road 268,Hangzhou 310029, China

Wende Liu The Key Laboratory of Biopesticide and Chemistry Biology, Min-istry of Education, Fujian Agriculture and Forestry University, Fuzhou 350002,P.R. China

Xiao-Hong Liu Biotechnology Institute, Zhejiang University, Kaixuan Road 268,Hangzhou 310029, China

Yan Liu Plant Breeding, Genetics and Biotechnology Division, International RiceResearch Institute, Manila, Philippines

Jian-Ping Lu Biotechnology Institute, Zhejiang University, Kaixuan Road 268,Hangzhou 310029, China

Guo-dong Lu The Key Laboratory of Biopesticide and Chemistry Biology, Min-istry of Education, Department of Plant Pathology & Microbiology, Texas A & MUniversity, College Station, USA 77843; College of Plant Protection, Fujian Agri-cultural and Forestry University, Fuzhou, China 350002

Erwina Lubis Indonesian Center for Rice Research (ICRR), Jalan Raya Ciapus25C, Muara, Bogor, West Java, Indonesia

Suzette Madamba Plant Breeding, Genetics and Biotechnology Division, Inter-national Rice Research Institute, Manila, Philippines

Cesar Martinez Centro Internacional de Agricultura Tropical, CIAT AA 6713Cali, Colombia

Hideo Matsumura Iwate Biotechnology Research Center, Kitakami, Japan

Shigeyuki Mayama Faculty of Agriculture, Kobe University, Kobe 657-8501,Japan

Thomas K. Mitchell Department of Plant Biology, Ohio State University, Colum-bus, OH, mitchell.815@osu.edu

xxvi Contributors

Orlando P. Morais Embrapa Rice and Beans, Caixa Postal 179, CEP 75375-000,Santo Antonio de Goias, GO, Brazil

J.B. Morel UMR BGPI INRA/CIRAD/Montpellier SupAgro, Campus Interna-tional de Baillarguet, TA A54/K, 34398 Montpellier, France, jbmorel@cirad.fr

Gloria Mosquera Department of Plant Pathology, Kansas State University,Manhattan, Kansas, 66506-5502, USA

Koji Nakagomi National Agricultural Research Center for TohokuRegion(NARCT), Morioka, Iwate Japan

Toshiki Nakamura National Agricultural Research Center for TohokuRegion(NARCT), Morioka, Iwate Japan

Hitoshi Nakayashiki Faculty of Agriculture, Kobe University, Kobe 657-8501,Japan

Naweed I. Naqvi Fungal Patho-Biology Group, Temasek Life SciencesLaboratory, 1 Research Link, National University of Singapore, Singapore 117604,naweed@tll.org.sg

D.C.S. Netto Centro de Ciencias Agrarias, Universidade Federal de Sao Carlos,FAPESP (The State of Sao Paulo Research Foundation) Scholars, P.O.Box 153, CEP13600-000 Araras, Sao Paulo, Brazil

Keita Nishide Laboratory of Plant Molecular Genetics, Nara Institute of Scienceand Technology, Ikoma, 630-0192 Japan

Jean-Loup Notteghem UMR BGPI INRA/CIRAD/Montpellier SupAgro,Campus International de Baillarguet, TA A54/K, 34398 Montpellier, France

In-Seok Oh National Institute of Crop Science, RDA, Suwon, Korea

Myung-Kyu Oh National Institute of Crop Science, RDA, Suwon 441-857, Korea

Yeon Yee Oh Center for Integrated Fungal Research, North Carolina State Univer-sity, Raleigh, NC

Jun Okuda Laboratory of Plant Molecular Genetics, Nara Institute of Science andTechnology, Ikoma, 630-0192 Japan

Isabelita Ona Plant Breeding, Genetics and Biotechnology Division, InternationalRice Research Institute, Manila, Philippines

Contributors xxvii

Chan Ho Park Department of Plant Pathology, The Ohio State University, USA

Anne S. Prabhu Embrapa Rice and Beans, Caixa Postal 179, CEP 75375-000,Santo Antonio de Goias, GO, Brazil, prabhu@cnpaf.embrapa.br

Dong-Soo Ra Plant Pathology Division, National Institute of Agricultural Scienceand Technology, RDA, Suwon, Korea

Ravikrishna Ramanujam Fungal Patho-Biology Group, Temasek Life SciencesLaboratory, 1 Research Link, National University of Singapore, Singapore 117604

Minoo Razee The School of Life Sciences, Fujian Agriculture and Forestry Uni-versity, Fuzhou 350002, P.R. China

Hee-Sool Rho Department of Pharmacology, Johns Hopkins School of Medicine,Baltimore, MD

Jae-Hwan Roh National Institute of Crop Science, RDA, Suwon 441-857, Korea

Hiromasa Saitoh Iwate Biotechnology Research Center, Kitakami, Japan

R.B.S. Sangar Indira Gandhi Agricultural University, Raipur, Chattisgarh

Santoso Indonesian Center for Rice Research (ICRR), Jalan Raya Ciapus 25C,Muara, Bogor, West Java, Indonesia; Research Institute for Rice, Subang, Indonesia

E. Seyran University of Arkansas, Division of Agriculture, Fayetteville, Arkansas72701, USA

Junjun Shang State Key Laboratory of Plant Genomics & National Plant GeneResearch Centre (Beijing), Institute of Genetics and Developmental Biology, Chi-nese Academy of Sciences, Beijing 100101, China; Current address: Department ofPlant Pathology, University of California, Riverside, CA 92521, USA

Yue Shang Department of Plant Pathology & Microbiology, Texas A & MUniversity, College Station, USA 77843

Matt Shenton Iwate Biotechnology Research Center, Kitakami, Japan

Ko Shimamoto Laboratory of Plant Molecular Genetics, Nara Institute of Scienceand Technology (NAIST), 8916-5 Takayama, Ikoma, 630-0192 Nara, Japan

C.P. Silva Centro de Ciencias Agrarias, Universidade Federal de Sao Carlos,FAPESP (The State of Sao Paulo Research Foundation) Scholars, P.O.Box 153, CEP13600-000 Araras, Sao Paulo, Brazil

xxviii Contributors

Gisele B. Silva Embrapa Rice and Beans, Caixa Postal 179, CEP 75375-000, SantoAntonio de Goias, GO, Brazil

Valacia L. Silva Lobo Embrapa Rice and Beans, Caixa Postal 179, CEP 75375-000, Santo Antonio de Goias, GO, Brazil

Carol Soderlund Arizona Genomics Computational Laboratory, BIO5 Institute,University of Arizona, Tucson, AZ 85721, USA, cari@agcol.arizona.edu

Pattavipha Songkumarn Department of Plant Pathology, The Ohio State Univer-sity, USA

A. Stabili Centro de Ciencias Agrarias, Universidade Federal de Sao Carlos,FAPESP (The State of Sao Paulo Research Foundation) Scholars, P.O.Box 153, CEP13600-000 Araras, Sao Paulo, Brazil

Jung-Pil Suh National Institute of Crop Science, RDA, Suwon 441-857, Korea

Suwarno Indonesian Center for Rice Research (ICRR), Jalan Raya Ciapus 25C,Muara, Bogor, West Java, Indonesia, balitpamuara@telkom.net

Yoshitaka Takano Laboratory of Plant Pathology, Kyoto University, Kyoto, Japan

Ryohei Terauchi Iwate Biotechnology Research Center, Kitakami, Japan, ter-auchi@ibrc.or.jp

Didier Tharreau CIRAD, UMR BGPI, Campus International de Baillarguet, TAA54/K, 34398 Montpellier, France, tharreau@cirad.fr

Yukio Tosa Faculty of Agriculture, Laboratory of Plant Pathology, Kobe Univer-sity, Kobe 657-8501, Japan

Tetsu Tsurushima Faculty of Business, Hannan University, Matsubara 580-8502,Japan, tsuru@hannan-u.ac.jp

A.S. Urashima Centro de Ciencias Agrarias, Universidade Federal de Sao Car-los, CEP 13600-000 Araras; Faculdade de Engenharia de Alimentos, Universi-dade de Campinas, Cidade Universitaria Zeferino Vaz, CEP 13083-970 Campinas,FAPESP (The State of Sao Paulo Research Foundation) Scholars, Sao Paulo, Brazil,alfredo@cca.ufscar.br

Dwinita Utami Biology Molecular Division, RIFCB, Bogor, Indonesia

Guadalupe Valdovinos-Ponce Department of Plant Pathology, Kansas State Uni-versity, Manhattan, Kansas, 66506-5502, USA

Contributors xxix

Barbara Valent Department of Plant Pathology, Kansas State University, Manhat-tan, Kansas, 66506-5502, USA, bvalent@ksu.edu

Mukund Variar Central Rainfed Upland Rice Research Station, Po Box 48, Haz-aribag 825 301, India, mukund.variar@gmail.com

R.C. Venu Department of Plant Pathology, The Ohio State University, USA

E. Vergne UMR BGPI INRA/CIRAD/Montpellier SupAgro, Campus Interna-tional de Baillarguet, TA A54/K, 34398 Montpellier, France

Fernando Correa Victoria Centro Internacional de Agricultura Tropical, CIATAA 6713 Cali, Colombia; RiceTec, Inc. PO Box 1305, Alvin, TX 77512, fcor-rea@ricetec.com

Guo-Liang Wang The Department of Plant Pathology, The Ohio State Uni-versity, Columbus, OH, 43210, USA; Crop Gene Engineering Key laboratory ofHunan Province, Hunan Agricultural University, Changsha, Hunan China, 410128,wang.620@osu.edu

Xueyan Wang USDA-ARS Dale Bumpers National Rice Research Center, Stuttgart,AR72160, USA

Zonghua Wang The Key Laboratory of Biopesticide and Chemistry Biology, TheSchool of Life Sciences, Ministry of Education, Fujian Agriculture and ForestryUniversity, Fuzhou 350002, P.R. China, zonghuaw@163.com

Joe Win Sainsbury Laboratory, Norwich, UK

Hanno Wolf Department of Plant Pathology & Microbiology, Texas A & M Uni-versity, College Station, USA 77843

Hann Ling Wong Laboratory of Plant Molecular Genetics, Nara Institute of Sci-ence and Technology (NAIST), 8916-5 Takayama, Ikoma, 630-0192 Nara, Japan,h-wong@bs.naist.jp

Jianli Wu China National Rice Research Institute, Hangzhou, China

Donghe Xu Japan International Research Center for Agricultural Sciences(JIRCAS), 1-1, Ohwashi, Tsukuba, Ibaraki 305-8686, Japan,xudh@jircas.affrc.go.jp

Jin-Rong Xu Department of Botany and Plant Pathology, Purdue University, WestLafayette, IN 47907; Departmetn of Plant Pathology, The Ohio State University,Columbus OH, jinrong@purdue.edu

xxx Contributors

Jichen Xu State Key Laboratory of Plant Genomics & National Plant GeneResearch Centre (Beijing), Institute of Genetics and Developmental Biology, Chi-nese Academy of Sciences, Beijing 100101, China

Jing Xu National Key Laboratory of Plant Molecular Genetics, Institute of PlantPhysiology and Ecology, Shanghai Institutes for Biological Sciences, GraduateSchool of Chinese Academy of Sciences, Chinese Academy of Sciences, Shanghai200032, China

Masayuki Yamaguchi National Agricultural Research Center for TohokuRegion(NARCT), Morioka, Iwate Japan

Chang-In Yang National Institute of Crop Science, RDA, Suwon 441-857, Korea

Sae-June Yang National Institute of Crop Science, RDA, Suwon 441-857, Korea

Yinong Yang Department of Plant Pathology and Program in Cell and MolecularBiology, University of Arkansas, Fayetteville, Arkansas 72701, USA; Departmentof Plant Pathology and Huck Institutes of Life Sciences, Pennsylvania State Univer-sity, University Park, Pennsylvania 16802, USA, yuy3@psu.edu

Mary Jeanie T. Yanoria International Rice Research Institute (IRRI). DAPO Box7777, Metro Manila, Philippines

Narifumi Yokogami National agricultural research center for Hokkaidoregion(NARCH)

Junichi Yonemaru National Agricultural Research Center for TohokuRegion(NARCT), Morioka, Iwate Japan

Kentaro Yoshida Iwate Biotechnology Research Center, Kitakami, Japan

Yunjia Xiang Institute of Plant Protection, Sichuan Academy of Agricultural Sci-ences, Jingjusilu Road 20, Chengdu 610066, China

Yunliang Peng Institute of Plant Protection, Sichuan Academy of Agricultural Sci-ences, Jingjusilu Road 20, Chengdu 610066, China, pengyunliang@yahoo.com.cn

Shaohong Zhang Guangdong Academy of Agricultural Sciences, Guangdong,China

Chuanzhi Zhao The School of Life Sciences, Fujian Agriculture and ForestryUniversity, Fuzhou 350002, P.R. China

Contributors xxxi

Xinhua Zhao Department of Botany and Plant Pathology, Purdue University, WestLafayette, IN 47907

Zhen Qin Plant Protection Station under Sichuan Provincial Ministry of Agricul-ture, Wuhouci Street 4, Chengdu 610041, China

Xiang-zi Zheng The School of Life Sciences, Fujian Agriculture and ForestryUniversity, Fuzhou 350002, P.R. China

Zhenyu Zhang Institute of Plant Protection, Sichuan Academy of AgriculturalSciences, Jingjusilu Road 20, Chengdu 610066, China

Bo Zhou The Department of Plant Pathology, Ohio State University, Columbus,OH, 43210, USA; Crop Gene Engineering Key laboratory of Hunan Province,Hunan Agricultural University, Changsha, Hunan China, 410128

Jie Zhou The Key Laboratory of Biopesticide and Chemistry Biology, The Schoolof Life Sciences, Ministry of Education, Fujian Agriculture and Forestry University,Fuzhou 350002, P.R. China

Xiaoying Zhou Department of Botany and Plant Pathology, Purdue University,West Lafayette, IN 47907

Xiangjun Zhou Department of Plant Pathology and Program in Cell and Molecu-lar Biology, University of Arkansas, Fayetteville, Arkansas 72701, USA

Heng Zhu Department of Pharmacology, Johns Hopkins School of Medicine,Baltimore, MD

Lihuang Zhu State Key Laboratory of Plant Genomics & National Plant GeneResearch Centre (Beijing), Institute of Genetics and Developmental Biology, Chi-nese Academy of Sciences, Beijing 100101, China, lhzhu@genetics.ac.cn

Xudong Zhu State Key Laboratory of Rice Biology, China National Rice ResearchInstitute, Hangzhou, 310006, China

Xiao Yuan Zhu Guangdong Academy of Agricultural Sciences, Guangdong,China

Current Status and Future Prospectsfor Research on Blast Resistance in Rice(Oryza sativa L.)

Gurdev S. Khush and K.K. Jena

Abstract Rice is the most important food security crop and staple food of half ofthe world population. Major increases in rice production occurred during the pastfour decades of last century as a result of adoption of green revolution technol-ogy. Demand for rice continues to increase as a result of population increase andimprovement in living standards particularly in Africa and Latin America. However,rate of increase of rice production has slowed down. It is estimated that we willhave to produce 30% more rice in 2030. For this purpose we need rice varieties withhigher yield potential and greater yield stability. Breakdown of blast resistance is themajor cause of yield instability in several rice growing areas. Efforts are underwayto develop rice varieties with durable blast resistance. More than 40 major genes aswell as QTL for blast resistance have been identified. Monogenic resistance to blastis less stable but varieties with pyramided monogenes or QTLs are durably resistant.Rice research should focus on identifying more durably resistant genes, tagging ofthese genes with molecular markers and pyramiding these genes or QTLs throughmolecular marker-aided selection. Candidate gene identification through rice func-tional genomics has great potential for developing more durably resistant varieties.

Keywords Monogenes · QTL · Resistance breakdown · Durable resistance · Genenomenclature

1 Introduction

Rice is the world’s most important food crop and a primary source of food for morethan half of the world’s population. More than 90% of the world’s rice is grownand consumed in Asia where 60% of the world’s people live. Rice accounts for

K.K. Jena (B)Senior Scientist, Plant Breeding, Genetics and Biotechnology Division, International RiceResearch Institute, C/o IRRI-Korea Office, National Institute of Crop Science, RDA, 209 SeodunDong, Suwon 441–857, Republic of Koreae-mail: k.jena@cgiar.org

G.-L. Wang, B. Valent (eds.), Advances in Genetics, Genomics and Control of Rice BlastDisease, DOI 10.1007/978-1-4020-9500-9 1,C© Springer Science+Business Media B.V. 2009

1

2 G.S. Khush and K.K. Jena

35–75% of the calories consumed by more than 3.0 billion Asians. Major increasesin rice production occurred during last four decades due to the adoption of greenrevolution technology. However, the rate of growth of rice production has sloweddown. Whereas rice production increased at the annual growth rate of 2.49% during1970–1990, the annual growth rate was 1.70% during 1990–2000 and only 1.21%during 2000–2006. World rice stocks are at the lowest level since 1974 and price ofrice is showing upward trend in the domestic and international markets.

The population of rice consumers is continuing to increase and demand for riceis also going up due to improved living standards particularly in Africa. Accordingto various estimates we will have to produce 30% more rice by 2030. To meet thischallenge we need rice varieties with higher yield potential and greater yield stabil-ity. Although yield potential of rice is 10 tons per hectare, farmers on the averageharvest about 5 tons per hectare from irrigated lands. This yield gap is due to thelosses caused by biotic and abiotic stresses. Among the biotic stresses blast diseaseis most important.

2 Rice Blast Problem and Yield Loss

Blast disease affects rice production in all rice growing regions. However, the yieldlosses are most serious in temperate regions and upland condition. Stresses causedby low temperature in temperate areas and moisture stress under upland conditionsincrease the vulnerability of the crop to blast disease attack. During the last 30 years,there were three outbreaks of blast in China; 1982–1985, 1992–1994, and 2001–2005. The average blast infected area was more than 3.8 million hectares in 1982–1985 with yield losses of several million tons (Sun et al. 1999). In 1993, yield lossof 1.1 million tons was recorded in Southern China alone. In recent years 5.7 millionhectares of rice fields were affected by the blast disease.

Blast disease is a major constraint in certain agro-ecological regions of India.Eastern India has the highest occurrence of blast followed by North and South.Sometimes yield losses reach as high as 50% in upland conditions (Widawskyand O’Toole 1990). Production oriented survey is organized by Directorate of RiceResearch, Hyderabad every year to document constraints to rice production in var-ious parts of the country. Multidisciplinary teams of scientists survey the crop inthe 125 districts of the country. About 15% of the districts surveyed during 2001–2005 had severe blast disease. Eastern India had the highest incidence of blast over12-year period. Hill districts of Himachal Pradesh, Uttaranchall, and Jammu andKashmir with cool climate generally have higher incidence of blast (Variar 2007).

The cool temperate climate of Japan is highly conducive to the multiplication andspread of the blast fungus during the rice growing season. Even though breeding forblast resistance has been emphasized in Japan since the beginning of last century anda number of genes for blast resistance have been introduced in Japanese varieties.Frequent breakdown of resistance results in significant yield losses ranging from 20–100% in some areas. In 1993 exceptionally cool weather resulted in blast epidemic.

Research on Blast Resistance in Rice (Oryza sativa L.) 3

Nation wide yield loss that year was 45.2% due to blast damage and sterility causedby low temperature and Japan was forced to import large quantities of rice fromabroad.

South Korea with temperate climate is another country where blast disease causesyield losses every year. During the blast epidemic of 1984–1985 almost 20% of therice area was seriously blasted.

In Indonesia 1.1 million hectares are planted to upland rice where blast is a seri-ous problem. Upland rice varieties become susceptible within one to two years oftheir cultivation. Yield losses will be as high as 100% during certain years. In 2004several hundred hectares of upland rice in Lampung province could not be harvesteddue to blast damage and yield losses up to 70% were recorded in upland areas ofWest Java (Sobrizal et al. 2007).

3 Genetics of Resistance

Studies on inheritance of resistance were first reported in 1922 by Sasaki followedby Takahashi (1965) and Yamasaki and Kiyosawa (1966). Up to 1960 the inheri-tance was studied without sufficient knowledge of the pathogenic specialization ofthe causal fungus, and it was quite difficult to obtain universal understanding of thesubject. Systematic studies were undertaken only after Goto (1965) established thedifferential system for blast fungus races in Japan.

Kiyosawa and his colleagues used seven Japanese strains of blast fungus forinvestigating the inheritance of resistance and identified 13 genes for resistance(Kiyosawa 1981). These were designated as Pi-a, Pi-i, Pi-ks , Pi-k, Pi-z, Pi-ta, Pi-ta2, Pi-zt Pi-k p, Pi-km, Pi-kn , Pi-b, and Pi-t. With the establishment of RiceGenetic Cooperative in 1985, rules for gene symbolization in rice were standard-ized and it was decided to assign gene symbols derived from names which describethe character modifications. The genes that are non-allelic but are indistinguishablefrom each other phenotypically, are designated by the same base letter but are differ-entiated by a number or a letter subscript separated by a hyphen (Kinoshita 1986).Since then several genes for resistance have been designated by base letter followedby a number subscript starting with Pi-1, Pi-2 (Mackill and Bonman 1992) and soon up to Pi-44.

During the third International Rice Genetics Symposium held at Manila on theOctober 16–19, 1995, it was decided to remove the hyphen between the base letterand subscript. Henceforth, the gene symbols have been written as Pia, Pita, Pik,Pi1, Pi2 etc.

During the past 25 years great advances have been made in studying the geneticsof resistance to the blast disease. Following conventional genetic analysis of iden-tified donors with resistance, availability of pure isolates of the blast pathogen anduse of advanced molecular analysis techniques, about 60 genes for resistance havebeen identified. These have been designated as Pi1, Pi2, Pi3, Pi4, Pi5, Pi6, Pi7, Pi9,Pi10 and Pi11 (Causse et al. 1994, Wang et al. 1994), Pia, Pib, Pik, Pit, Pita, Pita2,