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Arthur Kelman: Tribute and Remembrance

Luis Sequeira
Department of Plant Pathology, University of Wisconsin, Madison, Wisconsin 53706; email: lzs@plantpath.wisc.edu

Annu. Rev. Phytopathol. 2012.50:15-21. Downloaded from www.annualreviews.org by D'Youville College on 02/26/13. For personal use only.

Annu. Rev. Phytopathol. 2012. 50:1521 First published online as a Review in Advance on May 29, 2012 The Annual Review of Phytopathology is online at phyto.annualreviews.org This articles doi: 10.1146/annurev-phyto-010412-120339 Copyright c 2012 by Annual Reviews. All rights reserved 0066-4286/12/0908-0015$20.00

Keywords
bacterial plant pathogens, virulence, control of plant diseases, teaching plant pathology, professional societies

Abstract
With the death of Professor Arthur Kelman at age 90, the plant sciences, and particularly the eld of plant pathology, lost one of its most inuential and effective leaders. His long career involved important positions in the Departments of Plant Pathology at North Carolina State University (19491965) and the University of Wisconsin (19651989). Recognized not only for his achievements in research and his charismatic inuence on his numerous students, Arthur was, most of all, an extraordinary teacher. He and his students made signicant contributions in the elds of virulence, ecology, and control of bacterial plant pathogens. In addition, he was extremely active in service to professional societies, including the American Phytopathological Society, and a major force in the development of the International Society for Plant Pathology. In addition, he was an inuential member of the Council of the National Academy of Sciences, and, after retirement from the University of Wisconsin, he served for two years as Chief Scientist of the Competitive Grants Program of the US Department of Agriculture. He received numerous awards as a teacher and researcher, and felt an intense sense of duty to his profession. He was one of the most inuential plant pathologists of the twentieth century and will be remembered for his great intellect and his humanity.

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INTRODUCTION
I am pleased and honored to have an opportunity to write this memorial for Arthur Kelman, a truly distinguished scientist, an extraordinary leader, and close friend, who, more than anyone I can think of, guided plant pathology into the world of modern plant sciences, in the United States as well as internationally, during the past century. Plant pathology, as with other elds of science, continues to grow and improve because of the efforts of relatively few individuals who have had the intelligence as well as the personality to inuence the direction and impact of this particular area. Arthur Kelman was such a man. His imprint on plant pathology was rst established as a Distinguished Professor of plant pathology at North Carolina State University in Raleigh, NC, and later as chairman of the Department of Plant Pathology at the University of Wisconsin in Madison, WI. Beyond those signicant professional activities, Arthur became a primary supporter of (a) the change from volunteer to professional management of the American Phytopathological Society (APS), (b) the establishment of the International Society of Plant Pathology, and (c) basic agricultural research as chief scientist of the Competitive Grants Program of the US Department of Agriculture (USDA). I have written fairly recently (11) about Arthurs contributions to plant pathology, but the emphasis at that time was on anecdotes and other aspects of his lighter side. I have included some of those contributions in this paper. My apologies to those who feel that they have read some of this material before. Arthur was born in 1918 in Providence, RI, the youngest of a family that included eight children, all intensely active and entrepreneurial. He entered the University of Rhode Island in 1937, where he was exposed to plant pathology for the rst time. He found this eld so interesting that he initiated graduate work at NC State, but his career was interrupted by World War II. Because of his academic training and his ability to read and speak German, he was assigned to the Signal Intelligence Unit, working in North
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Africa, Sicily, and Italy, primarily with the purpose of decoding radio messages from the German command. He received a eld commission as a second lieutenant, but his work was interrupted when he was injured by shrapnel during the siege at Monte Casino in Italy. In 1946, after the war and after a short stint at the University of Wisconsin, where he was a student of J.C. Walker for a semester, Arthur returned to NC State to continue his career in plant pathology. A remarkably private person, Arthur never revealed to me why he left Madison. Returning to North Carolina, he met and married Helen Parker, a member of a distinguished local family and a teacher of biology at Meredith College in Raleigh. Helen, who passed away in 2011, was a very proper, sweet lady who embodied all the virtues that one associates with the quiet demeanor of the Southern lifestyle. Arthur was very much in love with her throughout his life, and he often had to curtail his competitive Yankee spirit in her presence. There were two children from this marriage, Philip, who is currently a musician in New York City, and a baby girl who died from a bacterial infection soon after birth. Arthur often remarked that his interest in bacteria probably stemmed from that unfortunate event. I was fortunate that my contact with Arthur extended for more than 50 years. Back in 1955, I supported his appointment as consultant for my employer (United Fruit Company); later, in 1960, I was a research associate in his laboratory at NC State. I was already a member of the Department of Plant Pathology at the University of Wisconsin when Arthur became chairman in 1964. After a period of 10 years as chairman, he remained in the department, and we continued to collaborate until his retirement in 1989. My admiration for his keen intellect and admirable ability to deal with all kinds of people never agged during that long period. He always seemed to know what do to or what to say at all occasions. Very guarded about his own private life, he had an extraordinary ability to get people to describe their problems and concerns to him, and he was always willing to help.

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Arthur became my close friend over the years, but he never allowed that close friendship to alter decisions that he had to make as chairman of the department. Perhaps his most outstanding quality was that he was always a fair person in all aspects of his relationship with staff, students, colleagues, and university administrators. Before he arrived at the University of Wisconsin, Arthur had enjoyed a remarkable career at NC State. He had become popular as an outstanding teacher of plant pathology as well as a devoted researcher of forest diseases. Severe local epidemics of bacterial wilt of tobacco that were caused by a bacterium (Pseudomonas solanacearum, now known as Ralstonia solanacearum) led him to prepare a comprehensive review of the literature on this important plant disease (4). Arthur indicated to me that his rst few years on the staff at NC State were so busy because of teaching responsibilities that he had no time to do laboratory research. He opted to search the literature, which resulted in the excellent review, published in 1953, entitled The Bacterial Wilt Caused by Pseudomonas solanacearum. That review, which is very useful even today, was typical of Arthurs approach: thorough, well-organized, rich in detailed descriptions of the literature, and excellent in his analysis of the signicance of the various contributions. It was that review that rst brought Arthur to my attention as I struggled in my rst job with attempts to deal with a serious epidemic of Moko disease (caused by a strain of the same bacterium) in the Coto River Valley of southwest Costa Rica. I was an employee of United Fruit Company, and at that time (19531960) the Coto River Valley was one of the most remote and isolated areas of the entire country of Costa Rica. In 1956, I attended a meeting of APS in Atlanta, where I listened to Arthur describe a simple means of preserving cultures of that bacterium (in distilled water), but, hard to believe, I was too shy to approach him and describe some of my own work. Yet, upon my return to Costa Rica, I managed to convince the research director of United Fruit Company in Boston that Arthur would be an excellent consultant.

Eventually, Arthur arrived at Coto 47, which was a collection of a few bungalows, a rather primitive laboratory, and little else in the midst of an oppressive jungle along the rail line that connects the town of Golto with the Panamanian border. Bananas were being planted in this new division, and Moko disease was already a big problem in the new farms. Arthur spent a couple of weeks with me at this unlikely location, and after numerous visits to the eld, he supported my notion that the source of the bacterium was the wild Heliconia species that grew everywhere. We made plans to study the various strains of the bacterium, their pathogenicity, their control, etc. Arthur was immensely helpful because of his profound knowledge of the literature. In addition, by that time Arthur had developed a rapid system for identication of the bacterium, based on addition of a dye, tetrazolium chloride (TZC), to an ordinary nutrient medium (5). The use of TZC medium, in which colonies of the bacterium have a very distinctive appearance, allowed me to survey many thousands of acres of bananas of the Golto Division for Moko disease and to design control procedures. In addition, Arthurs simple system for preserving the bacterium in distilled water allowed us to collect multiple strains of the bacterium, some capable of attacking bananas in addition to heliconias (1, 7). But it could not be all work. The research administrator at the time, Ira Hubbard, arranged for four of us to travel to Golto for a shing trip that included the use of the managers fancy boat, normally reserved for the big shots from Boston when they visited the division. The problem was that Arthur was not interested in outings or shing. A highly conservative person when it came to personal attire, he had trouble dispensing with his tie even in the oppressive heat of the banana plantations. When we arrived at the pier in Golto, most of us remained in the back of the boat, along with the shing equipment, beer, and food. But not Arthur; he selected a spot at the very front of the boat, his camera secure across his shoulder. He told me later that he did this because he did
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not feel safe in a small shing boat. Whether he was prescient or not, I cant say, but most of us were perfectly happy shing (no luck), eating, and drinking beer in back of the boat in the midst of a beautiful, sunny day. The captain chose a route that brought us close to the coast on our right and to a small island on our left that had a rocky promontory. That promontory prevented the captain from seeing a huge wave that was approaching us: a tsunami-like phenomenon probably born from some cataclysmic event far in the Pacic Ocean. By the time most of us and the captain realized what was happening, however, it was too late to try to turn the boat so as to front that gigantic wave. We were hit from the side. As the wave broke over the small island, all of us in back of the boat had just enough time to hang on to the nearest post. We were hit so hard that the boat nearly toppled over. When we recovered from that hit, the back of the boat was full of water, and we had lost every bit of shing equipment, the cameras, the beer, and the food. Immediately, we were all busy bailing out the water that threatened to sink the boat. We looked like a bunch of drenched rats but did not lose anyone, which was a miracle. I immediately searched for Arthur, who had been all alone in the front of the boat throughout the whole event. He had someow managed to hang on, and he kept his camera and did not get wet, which was quite unbelievable. All we could think was that as the back of the boat nearly sank from the weight of people and water, the front of the boat rose and was not hit as hard. I have described this event in some detail only to illustrate an important aspect of Arthurs personality. He was cool and collected throughout. Always generous and analytical, he refused to make a big deal of the tsunami event, and upon our return he thanked Ira Hubbard for an interesting trip. Over the next few years, I saw Arthur quite regularly in his role as consultant, and we collaborated extensively. He sent his graduate student, Charles Averre, to work with me for an entire semester. As a result, Charles and I provided the rst evidence of the distribution of
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different strains of a bacterial pathogen in the wild. I admired Arthurs sincerity as well as the personality that attracted people to him. He had charisma; people listened to him when he talked. He was sensitive to peoples problems and aspirations. On various occasions, he remarked that problems are caused quite often by people who lack antennae, a feature that allows insects to test the environment around them. He meant that you have to be careful not to offend others merely because you disagree with them. As our research projects progressed, Arthur suggested that I take leave to work with him in North Carolina. This offer, as it turned out, was quite providential. In late 1959, I had a serious disagreement with the vice president for research of United Fruit. As a result, he vowed to move me to the main laboratories in Honduras. I knew that I had no future with United Fruit as long as that person was in charge, and rather than to move to Honduras, I requested leave to work with Arthur at NC State. This was approved, and in January of 1960, I moved to Raleigh with my family. Arthur, his wife Helen, and his son Philip could not have been more gracious and hospitable. At the university, I was intensely happy doing research for over a year, going back to my early interest in hormone metabolism in diseased plants (8). One of Arthurs graduate students, Ellsworth Maine, had interviewed for a position at the University of Wisconsin in Madison and was prepared to accept it if offered to him. Unfortunately, he became ill and died a few months later, a victim of an aggressive form of leukemia. Arthur suggested my name as a possible alternative, and indeed, Glenn Pound arranged to interview me in January of 1961. It was not without some trepidation that I accepted the position when it was offered to me. My experience was with tropical plant diseases, and I worried about my familys reaction to the cold winters in Wisconsin. A couple of years after I joined the department, Glenn announced that he had agreed to become Dean of the College of Agriculture, and

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he established a committee that would search for his replacement. I could think of no one who would t that bill better than Arthur Kelman, and I indicated my strong support to the committee. Evidently, others agreed after Arthurs interview, and he was offered the position. He arrived at the department in July of 1965, a few months after the move to new quarters in Russell Laboratories. Thus, he inherited a strong tradition of excellent administrators (L.R. Jones, George Keitt, and Glenn Pound) and a group of strong senior professors ( J.C. Walker, Joyce Riker, and James Dickson) who frequently disagreed on almost any issue but who closed ranks when the department had to make important decisions. It did not take long for Arthur to establish his style of management. In contrast to Glenn Pound, who was direct and efcient but allowed little room for discussion, Arthur searched for everyones opinion when controversial issues came up for discussion at staff meetings. The fact is that Arthur had already approached everyone involved and knew precisely what was going to happen. There were never any surprises at staff meetings. He was calm and impartial. He knew exactly what to say on all occasions. He had a strong sense of humor, although he was in the habit of repeating the same stories many times over the years. Primarily, the staff appreciated his kind demeanor and his ability to prevent confrontations. His house became the center of numerous social interactions, particularly when visitors arrived in the department. He established high standards for the department and managed to convince the USDA to establish a Pioneer Research Center in our department, under the leadership of Richard Durbin. Arthurs idea was to have the ve members of the USDA group focus on resistance to a single disease. As it turned out, each member chose a separate disease problem and took off in different directions, much to Arthurs dismay. At Wisconsin, in addition to his extensive administrative and teaching duties, Arthur became interested in storage problems in potatoes and was helpful to growers in establishing the

conditions that would limit the damage done by soft-rotting bacteria (2, 9). In the 1970s, Arthur had to deal with a lot of turmoil at the university during the Vietnam War demonstrations. Teaching and research were affected. Some of our students joined the demonstrations, and Arthur dealt with them in a positive and generous manner. Although highly liberal in political outlook, Arthur managed to maintain a very impartial attitude in a department that was mostly conservative. Most impressive was Arthurs ability to teach the basic courses in plant pathology and to guide several graduate students to do research on storage diseases of potato, in spite of the demanding role of chairman. When he left NC State, he had already been named a Distinguished Professor, and it did not take long for the University of Wisconsin to bestow a similar honor. In 1976, he was elected to the National Academy of Sciences, where he was an inuential member of the council. He served the APS on numerous committees and eventually as president. He insisted on bringing professional help to run APS business, and when I complained openly that this would be too costly, he reminded me that I was talking out of ignorance. Indeed, the move to professional help turned out to be one of the wisest and most rewarding changes for APS. Arthur also spent time as president of the International Society for Plant Pathology. By the time he retired in 1989, Arthur (Figure 1) was one of the most widely respected professors at the University of Wisconsin and had received numerous awards over the course of his career. Those that deserve special mention are the Distingished Teacher Award and the William Neal Reynolds Distinguished Professor of Plant Pathology in 1961 at NC State. At Wisconsin, he was named the L.R. Jones Professor of Plant Pathology in 1975. In 1963, APS gave him its highest honor, the Award of Distinction. After he returned to NC State, Arthur received an Outstanding Alumnus Award from the College of Agriculture in 2000. At his retirement party, one of his sisters complained that Arthur was responsible for her
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Figure 1 Arthur Kelman. Photo courtesy of University of Wisconsin-Madison Department of Plant Pathology.

many hours of solitary connement when they were growing up because of Arthurs habit to appeal to their mother any time he was teased. Arthur was the youngest of a large family in Providence and the frequent target of his many brothers and sisters. As an adult, though, he maintained a deep affection for all members of his family. Arthurs remarkable inuence in the entire eld of plant pathology can be viewed with greater accuracy if one considers the state of knowledge of bacterial pathogens in the early 1950s (10). There was, at the time, only a handful of plant pathologists worldwide with an interest in fundamental knowledge of bacterial pathogens. Except for crown gall, caused by Agrobacterium tumefaciens, and several species of rhizobia (the cause of legume root nodules), most plant-pathogenic bacteria seemed to inspire no interest among plant pathologists. An important revolution, led by Arthur in the
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1950s, transformed that initial lack of interest. Consider the fact that nowadays several hundred research papers on the basic sciences dealing with the interactions of bacteria and their plant hosts are published each year. A quick perusal of recent publications, such as Phytopathology and Molecular Plant-Microbe Interactions (MPMI), reveals that fully one-third of the monthly research contributions deal with bacterial-plant host interactions. It might be useful to consider how that change came about. By the middle of the twentieth century, most plant pathologists interested in host-pathogen interactions had followed the traditions established by Anton de Bary and H. Marshall Ward and were concerned with the nature of the enzymes that degrade pectic substances in plant cell walls (10). The host was considered a rather inanimate source of substrata for the cell wall degrading enzymes of plant pathogens. There were indications, however, that host responses to infection determined the fate of the interaction. It was Ward, after all, who described the death of the cells adjacent to the infection site in grasses resistant to rust fungi. The term hypersensitive reaction (HR) was later introduced to describe this type of interaction, but its significance remained obscure until Harold Flor described gene-for-gene interactions in rust diseases of ax. For bacterial diseases of plants, it was Arthur Kelman who, in the 1950s, began to establish the connection between the production of extracellular polysaccharides (EPS), motility, and virulence in R. solanacearum (3, 6). That became the fountainhead of research on the relationship of EPS to induction of the HR in tobacco and other hosts for many other plant pathologists. That was also the beginning of an extensive list of publications by numerous plant pathologists on the molecular genetics of bacterialplant host interactions that continues unabated to this day. Arthurs retirement from the University of Wisconsin in 1989 had a profound impact on the Department of Plant Pathology in ways that are probably not fully recognized by my colleagues. In recent years, the department has been unable to replace many of the positions of

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retired members and has become more divisive. Arthur maintained the tradition of respect for the past and of good relations among staff, even when opinions differed. He was able to garner support for departmental programs in spite of cutbacks in the university budget. Special mention should be made of Arthurs special contributions as a member of the Council of the National Academy of Sciences. He realized that agricultural sciences were receiving short shrift when compared with astronomy, physics, mathematics, etc. Arthur was instrumental in creating a separate board on
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agriculture that over the years was highly instrumental in recognizing the basic science that is contributed by researchers on agriculture. After retirement, Arthur was head of the Competitive Grants Program of the USDA for two years. That program was, and still is (under a different rubric), one of the major sources of support for basic research in agriculture. That was a tting end to a long career dedicated to the advancement of the plant sciences. That career remains an important legacy for all young scientists who are considering a life in plant pathology.

DISCLOSURE STATEMENT
The author is not aware of any afliations, memberships, funding, or nancial holdings that might be perceived as affecting the objectivity of this review. LITERATURE CITED
1. Buddenhagen I, Kelman A. 1964. Biological and physiological aspects of bacterial wilt caused by Pseudomonas solanacearum. Annu. Rev. Phytopathol. 2:20330 2. Cuppels D, Kelman A. 1973. Evaluation of media for isolation of soft-rot bacteria from soil and plant tissue. Phytopathology 64:46875 3. Husain A, Kelman A. 1958. Relation of slime production to mechanism of wilting and pathogenicity in Pseudomonas solanacearum. Phytopathology 48:15565 4. Kelman A. 1953. The bacterial wilt caused by Pseudomonas solanacearum. N.C. Agric. Exp. Stn. Tech. Bull. 99. 194 pp. 5. Kelman A. 1954. The relationship of pathogenicity of Pseudomonas solanacearum to colony appearance on a tetrazolium medium. Phytopathology 44:69395 6. Kelman A, Hruschka. 1973. The role of motility and aerotaxis in the selective increase of avirulent bacteria in still broth cultures of Pseudomonas solanacearum. J. Gen. Microbiol. 76:17788 7. Kelman A, Person LH. 1961. Strains of Pseudomonas solanacearum differing in pathogenicity to tobacco and peanut. Phytopathology 51:15861 8. Kelman A, Sequeira L. 1965. Root-to-root spread of Pseudomonas solanacearum. Phytopathology 55:3049 9. Perombelon MCM, Kelman A. 1980. Ecology of the soft rot erwinias. Annu. Rev. Phytopathol. 18:36187 10. Sequeira L. 2001. Delivery of pathogen signals: historical approach. In Delivery and Perception of Pathogen Signals in Plants, ed. NT Keen, S Mayama, JE Leach, S. Tsuyama, pp. 1224. St. Paul, MN: APS Press 11. Sequeira L. 2010. Arthur Kelman: a man for all seasons. In And One Hand on the Bench, ed. JH Andrews, pp. 4852. Madison, WI: Univ. Wis. Press

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Contents
An Ideal Job Kurt J. Leonard p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 1
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Annual Review of Phytopathology Volume 50, 2012

Arthur Kelman: Tribute and Remembrance Luis Sequeira p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 15 Stagonospora nodorum: From Pathology to Genomics and Host Resistance Richard P. Oliver, Timothy L. Friesen, Justin D. Faris, and Peter S. Solomon p p p p p p p p p p 23 Apple Replant Disease: Role of Microbial Ecology in Cause and Control Mark Mazzola and Luisa M. Manici p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 45 Pathogenomics of the Ralstonia solanacearum Species Complex St ephane Genin and Timothy P. Denny p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 67 The Genomics of Obligate (and Nonobligate) Biotrophs Pietro D. Spanu p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 91 Genome-Enabled Perspectives on the Composition, Evolution, and Expression of Virulence Determinants in Bacterial Plant Pathogens Magdalen Lindeberg p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 111 Suppressive Composts: Microbial Ecology Links Between Abiotic Environments and Healthy Plants Yitzhak Hadar and Kalliope K. Papadopoulou p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 133 Plant Defense Compounds: Systems Approaches to Metabolic Analysis Daniel J. Kliebenstein p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 155 Role of Nematode Peptides and Other Small Molecules in Plant Parasitism Melissa G. Mitchum, Xiaohong Wang, Jianying Wang, and Eric L. Davis p p p p p p p p p p p p 175 New Grower-Friendly Methods for Plant Pathogen Monitoring Solke H. De Boer and Mar a M. L opez p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 197 Somatic Hybridization in the Uredinales Robert F. Park and Colin R. Wellings p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 219
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Interrelationships of Food Safety and Plant Pathology: The Life Cycle of Human Pathogens on Plants Jeri D. Barak and Brenda K. Schroeder p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 241 Plant Immunity to Necrotrophs Tesfaye Mengiste p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 267 Mechanisms and Evolution of Virulence in Oomycetes Rays H.Y. Jiang and Brett M. Tyler p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 295 Variation and Selection of Quantitative Traits in Plant Pathogens Christian Lannou p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 319
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Gall Midges (Hessian Flies) as Plant Pathogens Jeff J. Stuart, Ming-Shun Chen, Richard Shukle, and Marion O. Harris p p p p p p p p p p p p p p 339 Phytophthora Beyond Agriculture Everett M. Hansen, Paul W. Reeser, and Wendy Sutton p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 359 Landscape Epidemiology of Emerging Infectious Diseases in Natural and Human-Altered Ecosystems Ross K. Meentemeyer, Sarah E. Haas, and Tom as V aclav k p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 379 Diversity and Natural Functions of Antibiotics Produced by Benecial and Plant Pathogenic Bacteria Jos M. Raaijmakers and Mark Mazzola p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 403 The Role of Secretion Systems and Small Molecules in Soft-Rot Enterobacteriaceae Pathogenicity Amy Charkowski, Carlos Blanco, Guy Condemine, Dominique Expert, Thierry Franza, Christopher Hayes, Nicole Hugouvieux-Cotte-Pattat, Emilia L opez Solanilla, David Low, Lucy Moleleki, Minna Pirhonen, Andrew Pitman, Nicole Perna, Sylvie Reverchon, Pablo Rodr guez Palenzuela, Michael San Francisco, Ian Toth, Shinji Tsuyumu, Jacquie van der Waals, Jan van der Wolf, Fr ed erique Van Gijsegem, Ching-Hong Yang, and Iris Yedidia p p p p p p p p p p p p p p p p p p p p p p 425 Receptor Kinase Signaling Pathways in Plant-Microbe Interactions Meritxell Antol n-Llovera, Martina K. Ried, Andreas Binder, and Martin Parniske p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 451 Fire Blight: Applied Genomic Insights of the Pathogen and Host Mickael Malnoy, Stefan Martens, John L. Norelli, Marie-Anne Barny, George W. Sundin, Theo H.M. Smits, and Brion Duffy p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 475 Errata An online log of corrections to Annual Review of Phytopathology articles may be found at http://phyto.annualreviews.org/
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