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Physiological Plant Pathology PDF

908 Pages·1976·29.767 MB·English
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Encyclopedia of Plant Physiology New Series Volume 4 Editors A. Pirson, Gottingen M. H. Zimmermann, Harvard Physiological Plant Pathology Edited by R. Hei tefuss and P. H. Williams Contributors J.R.Aist P.J.Allen H.G.Basham D.FBateman J. M. Daly H. M. Dekhuijzen V H. Dropkin J. M. Duniway A. H. Ellingboe F Fric W H. Fuchs D. G. Gilchrist R. N. Goodman R. Heitefuss E. C. Hislop D. S. Ingram T. Kosuge J. A. Kuc B. B. Lippincott J. A. Lippincott K.Maramorosch J. E. Mitchell G. F Pegg D. Pitt K.Rudolph R. P. Scheffer E. SchlOsser F Schonbeck K. J. Scott R. C. Staples 1. Uritani H. E. Wheeler G. Wolf Z. Yaniv C. E. Yarwood G. A. De Zoeten Springer-Verlag Berlin Heidelberg New York 1976 With 92 Figures ISBN- 13: 978-3-642-66281-2 e-ISBN-13: 978-3-642-66279-9 DOl: 10.1007/978-3-642-66279-9 Library of Congress Cataloging in Publication Data. Main entry under title: Physiological plant pathology. (Encyclope dia of plant physiology: v. 4) Includes bibliographical references and indexes. 1. Plant diseases. 2. Plant physiology. I. Heitefuss. Rudolf. II. Williams, Paul. H., 1938 - .SB731.P53. 581.2.76-3724. This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under § 54 of the German Copyright Law where copies are made for other than private use, a fee is payable to the publisher, the amount of the fee to be determined by agreement with the publisher. by Springer-Verlag Berlin· Heidelberg 1976 Softcover reprint of the hardcover 1st edition 1976 The use of registered names, trademarks, etc. in this pUblication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Preface Plant pathology embraces all aspects of biological and scientific activity which are concerned with understanding the complex phenomena of diseases in plants. Physiological plant pathology represents those specialities within plant pathology which focus on the physiological and biochemical activities of pathogens and on the response of host plant tissues. Today there is an increasing recognition on the part of the scientific agri cultural community that only through a deeper and more fundamental under standing of all the interacting components of the agricultural biota can we expect to improve our capabilities of feeding an expanding world population. It is in this context that physiological plant pathology has assumed new significance within the broader field of plant pathology. No longer are studies on the biochemistry and physiology of pathogens and pathogenesis merely isolated academic exercises; rather, a substantial coherent body of knowledge is accumulating upon which our understanding of the process of disease developmen t and host resistance is being founded. It is from these foundations of knowledge that ultimately new insights into the control of plant diseases may be expected to grow. It seems appropriate, therefore, that at regular intervals those involved in the various subspecialities encompassing the broadest aspects of physiological plant pathology reassess the contributions within the particular specialities in the light of new knowledge and technologies for the purpose of articulating new and productive directions for the future. It has become increasingly apparent in all areas of biology that a full under standing of any particular phenomenon lies at the interfaces of knowledge from diverse disciplines. Thus, only by bringing together the insights of disciplines such as genetics, physiology, cytology, biochemistry and biophysics will the complexities and significance of host-pathogen relations be fully understood. With the publication of this volume, we hope to have provided researchers and students with the opportunity of assessing the state of our knowledge in diverse aspects of physiological plant pathology, ranging from the physiology and ultrastructural cytology of various pathogens to the biochemistry of pathogenesis and resistance. From throughout the world we have selected active researchers and asked them to review the past, to evaluate the present knowledge and to assess the future potential for their respective disciplines. We have selected two authors with especially long and thorough experience in the field of physiological plant pathology to introduce the volume. In a historical chapter we hope to provide the reader with a view of the beginnings and philo sophical roots from which modern physiological plant pathology has grown. In the following chapter, some of the principal aspects of host-pathogen interactions VI Preface in need of modem biochemical technology and analytical objectivity are critically evaluated. We have avoided dwelling on definitions and terminology in the intfoductory chapters but instead refer to the phytopathological paper No. 17 (1973) of the Commonwealth Mycological Institute as "a guide to the use of terms in plant pathology". The main part of the volume is structured with the realization that host-pathogen interactions progress through a series of stages before and during pathogenesis which are ultimately expressed as susceptibility or resistance of the plant. Before the parasite enters the host, physiological events occur which may be decisive in determining the ultimate success of the attack. Normally, these activities take place in the rhizosphere or phyllosphere and the relationship of the chemical and physical environments of these regions may be of great importance to the regulation of germination and penetration. Although most of the emphasis in this volume is on the reactions of higher plants with fungi and bacteria; viruses, nematodes and mycoplasm-like organisms have also been included. The specialist in these latter fields may miss some important aspects, as the scope of the volume has not permitted complete coverage. Our intention has been to focus on various similarities and differences within the different groups and in their reactions with the host. This should be especially evident in the section on physiology and cytology of penetration and establishment, an area in which the progress in knowledge depends primarily on the careful investigation and interpretation of structure and function. 1\ study of pathogenesis involves not only an understanding of the alteration in normal plant metabolism, water balance, and growth responses, but also a knowledge of those factors used by pathogens to aid or assist in the attack and colonization of host tissues. A number of chapters, therefore, are directed to the activities of toxins, enzymes, and growth regulators as factors involved in patho genesis and to alterations in metabolic pathways, water balance, and growth responses in the diseased plant. Of great interest today in studies of physiological plant pathology are the events which result in resistance to pathogens or in manifestations of specificity in particular combinations of host and pathogen genotypes. The chapters on pre formed and induced determinants of resistance and specificity review major research in these areas. Strongly complementing these discussions on resistance are chapters on the genetic basis for specificity and the role of metabolic regulation in the infected host tissue. Any consideration of the diseased plant must take into account possible factors which may have had an effect in predisposing that plant to disease. The influence of predisposition is often neglected by pathologists viewing diseases under conditions designed for physiological studies. For this reason, we thought it particularly important to include a chapter on predisposing factors in plant disease. Technological advances in the past 1O~15 years have now permitted the routine cultivation of a number of biotrophic parasites in axenic and monoxenic culture on host callus. The implications of these methodologies to cytological, physiological, and biochemical analysis of obligate parasitism are very great and it seemed appropriate to review the status of these emerging specialities. Outside the scope of this volume are the diseases or disorders of plants caused by non-parasitic agents. Therefore the physiology of disorders due to deficiency, excess Preface VII or imbalance of elements essential to plant growth has not been treated. It is hoped that these problems will be included in a later volume of this series. The same applies to injury of plants by phytotoxic air pollutants and other chemical agents. Just as it is expected that a knowledge of the physiological processes of disease in plants can be gained only after understanding the normal physiology in the plant, so it is appropriate to include in this New Series of the Encyclopedia of Plant Physiology a volume of Physiological Plant Pathology. Indeed, in such a setting as this series, the study of disease processes in plants may not only receive closer scrutiny by plant physiologists and biochemists, but physiological plant pathologists may be directed to new insights of the plant which provide solutions to their particular problems. Our thanks are due to the authors of this volume for their contributions and for their patience during the preparation of this book, which has taken longer than we had originally anticipated. We thank Professor Dr. W.H. FUCHS and Professor Dr. A. PIRSON, both in G6ttingen, for valuable suggestions on the outline and scope of this volume. Further thanks are extended to Mrs. KAUNA TH for her technical assistance. We appreciate the assistance by various members of the Springer-Verlag for the final preparation of the book. G6ttingen and Madison, R. HEITEFUSS May 1976 P.H. WILLIAMS Contents 1. General 1.1 History of Physiological Plant Pathology W.H. FUCHS ........... . 1.2 Some Aspects of Host-Pathogen Interactions 1.M. DALY (With 1 Figure) 1. Introduction . . . . . . 27 2. The Response of the Host 27 2.1 Resistance . . . . . 28 2.2 Susceptibility . . . . 30 2.3 Economic Damage and Tolerance to Disease 30 2.4 Symptom Expression and Spurious Effects . 31 3. Pathogenicity and Virulence . . . . . . . . . 32 4. The Influence of Disease in the Design of Experimental Procedures 34 5. The Control of Host-Pathogen Interactions. 35 5.1 Genetic Control. . . . . . . . 35 5.2 Flor's Gene-for-gene Hypothesis. . . . 36 5.3 Metabolic Control. . . . . . . . . . 37 5.4 Modes of Inheritance and Induced Resistance versus Induced Susceptibility 39 5.5 Recognition Phenomena in Specificity . . . . . . 40 5.6 The Influence of Environment. . . . . . . . . . 41 6. Special Topics . . . . . . . . . . . . . . . . . . 41 6.1 Quantitative Measures of Disease and of Pathogens 41 6.2 Genetic Control of Experimental Material 43 6.3 Model Systems . . . 44 6.4 Challenge Inoculations 45 References . . . . . . . . 47 2. Spore Germination and Its Regulation 2.1 Control of Spore Germination and Infection Structure Formation in the Fungi P.l. ALLEN 1. Introduction . . . . . . . . . . . . . . . . . . 51 2. Self-Inhibition and Stimulation of Gennination . . . 54 2.1 Gennination Inhibitors from the Parent Colony . 54 2.2 Gennination Inhibitors in Spores . . . . . . . 55 2.3 Self-Stimulation of Gennination. . . . . . . . 66 3. Inhibitors and Stimulants of Gennination in Soil. .. 68 3.1 Soil Fungistasis . . . . . . . . . . . . . . . 68 3.2 Stimulants in Soil . . . . . . . . . . . . . . 71 4. Stimulants and Inhibitors of Gennination at Plant Surfaces 73 4.1 Inhibitors 73 4.2 Stimulants . . . . . . . . . . . . . . . . . . . . 75 x Contents 5. Fonnation of Infection Structures. . . . . . . . . . . 76 5.1 Genetic Basis of Infection Structure Fonnation . . . 76 5.2 Experimental Studies on Infection Structure Fonnation 76 6. Conclusions 77 References . . . . . . . . . . . . . . . . . . . . . 78 2.2 Protein and Nucleic Acid Metabolism during Germination R.C. STAPLES and Z. Y ANIV (With 2 Figures) 1. Introduction . . . . . . . 86 2. Facultative Parasites 86 2.1 Gennination of Conidia 86 2.2 Gennination of Zoospores 93 2.3 Gennination of Basidiospores 95 3. Obligate Parasites. . . . . . . 96 3.1 Introduction ...... . 96 3.2 Changes in Proteins and Nucleic Acids during Gennination 96 3.3 Changes in Ribosome Activity during Spore Gennination 97 3.4 Differentiation 98 3.5 Conclusions. 99 References . . . . . 101 2.3 The Effect of Roots on the Activity of Soil-borne Plant Pathogens J .E. MITCHELL 1. Introduction . . . . . . 104 2. The Pathogen in the Soil . 105 3. The Root-Soil Interface 106 3.1 The Rhizosphere ... 106 3.2 The Rhizoplane . . . 107 4. The Mechanism of the Root Effect 108 4.1 Root Tissue Debris . . . . 108 4.2 Root Exudates . . . . . . . 108 4.3 Oxygen and Carbon Dioxide . 110 4.4 Volatile Emanations from Roots. 111 4.5 Root Surface Effects . . . . . . III 5. Methods of StUdying Response of Pathogens to Roots 112 6. Behavior of Specific Pathogens 113 7. Discussion 122 References . . . . . . . . . . 123 3. Cytology and Physiology of Penetration and Establishment 3.1 Cytology of Virus Infection and Virus Transport G.A. DE ZOETEN (With 8 Figures) 1. Introduction . . . 129 2. Methods of Study. . . . . . 130 3. Infection. . . . . . . . . . 132 4. Cytology of Virus Multiplication 136 5. Virus Aggregation, Anomalous Inclusions and Structural Changes in Infected Cells 141 6. Transport . 143 7. Conclusions 145 References . . 146 3.2 Plant Mycoplasma Diseases K. MARAMOROSCH (With 1 Figure) 1. Introduction . . . . . . . 150 2. Historical Background. . . 150 3. Tenninology and Definitions 152 Contents XI 4. Methodology. . . . . . . . . 154 5. Effects of MLO on Plants . . . 159 6. Effects of MLO on Insect Vectors 161 7. Chemotherapy and Heat Therapy 162 8. Unsolved Problems . . . 164 9. Summary and Conclusions . . . 165 References . . . . . . . . . . . 166 3.3 Physiological and Cytological Aspects of the Bacterial Infection Process R.N. GOODMAN (With 5 Figures) I. Introduction . . . . . . . . . . . . 172 2. Pre-penetration . . . . . . . . . . . 172 2.1 Motility, Aerotaxis and Chemotaxis 172 2.2 Predisposing Environmental Features. 174 2.3 Morphological Barriers to Infection 179 3. Penetration. . . . . 180 3.1 Natural Openings . . 180 3.2 Wounds . . . . . . 183 4. Growth of the Bacteria in vivo 186 4.1 Inoculum Size Required to Establish an Infection 186 4.2 The Sites of Bacterial Multiplication. . . . . . 186 4.3 Bacterial Proliferation in Connection with the Hypersensitive Reaction 190 4.4 Growth of Saprophytic Bacteria in Plant Tissue 190 5. Microbiological Antagonism and Commensalism 191 6. Summing-up 192 References . . . . . . . . . . . . . . . . 193 3.4 Cytology of Penetration and Infection - Fungi 1.R. AIST (With 12 Figures) I. Introduction . . 197 2. The Use of Modem Methodology 197 3. Primary Penetration. . . . . . 201 3.1 Primary Penetration Types . 201 3.2 The Mechanism(s) of Primary Penetration 206 4. Infection. . . . . . . . . . . . . . . . . 209 4.1 Specialized Structures Associated with Infection 209 4.2 Effects of Infection on Susceptible Host Cytology 213 5. Relating Structure to Function 215 References . . . . . . . . . . . . . . . . . . . . 216 3.5 Nematode Parasites of Plants, Their Ecology and the Process of Infection V.H. DROPKIN (With 3 Figures) 1. Introduction . . . . . . . . . . . . . . . 222 2. Physiology of Nematodes outside Their Hosts 223 3. Attraction and Penetration 229 3.1 Attraction . . . . . . 229 3.2 Feeding Behavior . . . 230 3.3 Wall-degrading Enzymes 232 4. Nutrition . . . . . . 232 4.1 Cultures . . . . . 232 4.2 Sex Determination. 233 4.3 Growth Rates. . . 234 4.4 Mineral Nutrition of Host 234 5. Cyto- and Histopathology . . . . . . 234 6. Biochemical Changes in Infected Tissues 240 7. Remarks on Breeding for Resistance 241 References . . . . . . . . . . . . . . 242 XII Contents 4. Forces by Which the Pathogen Attacks the Host Plant 4.1 Host-specific Toxins in Relation to Pathogenesis and Disease Resistance R.P. SCHEFFER (With 1 Figure) 1. Introduction . . . . . . . . . . . . . .247 2. The Agronomic Background . . . . . . 248 3. Development and Use of Toxin Bioassays 250 4. Genetic Studies of the Toxin Problem . . 254 5. Isolation and Characterization of Toxins . 255 6. Comparative Effects of the Fungi and Their Toxins on Susceptible Cells and Tissues . . . . . . . . . . . . . . 258 7. The Resistant Cell in Relation to Toxin 264 8. Summary 265 References . . . . . . . . . . . . . . 265 4.2 Non-specific Toxins K. RUDOLPH (With I Figure) I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . 270 2. Definitions. . . . . . . . . . . . . . . . . . . . . . . . . 271 3. Production of Phytotoxic Compounds in vitro by Plant Pathogens 272 4. Assay Systems for Phytotoxicity. . . . 274 5. Purification of Toxins . . . . . . . . 276 6. Lowest Effective Concentration or Doses 277 7. Chemistry of Phytotoxic Compounds. . 277 8. Mode of Action of Phytotoxic Compounds 283 9. Toxins in Relation to Pathogenesis . . 289 10. Biosynthesis and Degradation of Toxins . 294 11. Uses of Phytotoxic Compounds 295 12. Phytopathogens Which Produce Phytotoxic Compounds 296 13. Concluding Remarks 301 References. . . . . . . . . . . . . . . . . . . . . . 301 4.3 Degradation of Plant Cell Walls and Membranes by Microbial Enzymes D.F. BATEMAN and H.G. BASHAM (With 6 Figures) 1. Introduction . . . . . . . . . . . . . . . . . 316 2. Structural Components . . . . . . . . . . . . 316 3. Enzymes Which Degrade Plant Structural Elements 321 3.1 Cell Wall Degradation . . . . . . . . . . . . 321 3.2 Membrane Degradation . . . . . . . . . . . 327 4. Relationships of Enzymatic Alterations of Structural Constituents to Pathogenesis 329 4.1 Alterations of Cell Wall Constituents . 330 4.2 Enzymatic Degradation of Membranes. 342 5. Summary and Additional Considerations. 344 References . . . . . . . . . . . . . . . . 345 4.4 Morphogenic Determinants as Exemplified by the Crown-gall Disease 1.A. LIPPINCOTT and B.B. LIPPINCOTT (With I Figure) 1. Introduction . . . . . . . . . . . . . . . . 356 1.1 Historical Background and Related Diseases 356 1.2 Gall Morphology and Structure . . . . . . 357 1.3 General Concepts . . . . . . . . . . .. 357 2. The Causal Agent . . . . . . . . . . . . . . 360 2.1 Taxonomic Relationships of the Agrobacteria 361 2.2 Agrobacterium Genetics . . . . . . . . . 362

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