Progress in Experimental Tumor Research
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1 Plant Tumor Research
2 Progress in Experimental Tumor Research voι, ιs Editor: F. HOMBTJRGER, Cambridge, Mass. S. Karger Basel München Paris London New York Sydney
3 Plant Tumor Research Guest Editor: Α. C. BRAUN, New York, N.Y. Contributors: R. E. BEARDSLEY, Bronx, Ν.Υ. ; L. Μ. BLACK, Urbana, Ill.; Α. C. ΒR UΝ, New York, Ν.Υ. ; F. MEWS, jr., Princeton, N.J. ; H.H. SωΤκ, Upton, N.Y.; H.N. Woon, New York, N.Y. With 32 figures and 2 tables S. Karger. Basel München Paris London New York Sydney 1972
4 Progress in Experimental Tumor Research Vol. 1: XVΙ p., 166 fig., 44 tab., 1 cpl., 1960 Vol. 2: XIV p., 220 fig., 48 tab., 1961 Vol. 3: XVI p., 158 fig., 20 tab., 1963 Vol. 4: XII p., 169 fig., 41 tab., 1964 Vol. 5: XII p., 85 fig., 48 tab., 1964 Vol. 6: XII p., 45 fig., 34 tab., 1965 Vol. 7: VIII p., 43 fig., 26 tab., 1965 Vol. 8: VI p., 93 fig., 36 tab., 1966 Vol. 9: X p., 85 fig., 25 tab., 1967 Vo1.10: X p., 40 fig., 15 tab., 1968 Vo1.11: XII ρ., 92 fig., 73 tab., 1 cpl., 1969 Vo1.12: VIII p., 11 fig., 10 tab., 1969 Vo1.13 : Immunological Aspects of Neoplasia, X p., 21 fig., 6 tab., 1970 Vo1.14: Inhibition of Carcinogenesis, X p., 34 fig., 20 tab., 1971 S. Karger Basel München Paris London New York Sydney Arnold-Böcklin-Strasse 25, CH-4000 Basel 11 (Switzerland) All rights, including that of translation into other languages, reserved. Photomechanic reproduction (photocopy, microcopy) of this book or part of it without special permission of the publishers is Copyright 1972 by S. Karger AG, Verlag für Medizin und Naturwissenschaften, Basel Printed in Switzerland by Basler Druck- und Verlagsanstalt AG, Basel Blocks: Steiner & Co., Basel
5 Contents Series Editor's Foreword Guest Editor's Foreword VIII IX The Inception Phase in the Crown Gall Disease R. E. BEARDSLEY I. Introduction 1 A. Evidence for Bacteria-Free Tumor Tissue 2 B. The Concept of the Tumor-Inducing Principle 6 II. The Role of Wounding in Tumorigenesis 10 A. The Portal of Entry Hypothesis 12 B. Tumorigenic Competence Competence and Wound Healing Competence and Progression Competence and Morphogenetic Regulants Competence and Association-Induction Phenomena Populations of Competent Cells 35 III. Conversion 38 A. Conversion and the Reduced State 40 B. Theories of Conversion 45 C. The Two-Stage Hypothesis of Conversion 58 IV. Conclusions 61 References 62 The Development of a Capacity for Autonomous Growth of the Crown Gall Tumor Cell H. N. WOOD I. The Inception Phase 78 II. The Developmental Phase 79
6 Contents VI III. The Regulatory Function of Membranes in Crown Gall Tumorigenesis IV. The Control of Cell Division and the Natural Growth Regulators in Normal and Plant Tumor Cells 86 V. Summary and Conclusions 89 References 90 Stability of the Tumor Phenotype in Crown Gall Tumors of Tobacco F. Μεπνs, jr. I. Introduction 93 II. Reversibility of the Tumor Phenotype in Tobacco 94 III. Stability of the Teratoma Phenotype in Culture 95 IV Chemically Induced Changes in Teratoma Phenotype 98 V. Possible Mechanisms for Tumor Stability 101 VI. Summary and Conclusions 105 References 107 Plant Tumors of Viral Origin L. M. BLACK I. Introduction 110 II. Experimental Findings 116 A. Tumor Variations Related to the Heredity of the Individual 116 B. The Role of Wounds in the Development of Tumors in Virus-Infected Plants 119 C. The Possible Role of Hormones 124 D. Recovery 124 E. Culture of Plant Tissues in vitro 125 F. The Complex Biological Relationships between the Virus and Its Insect Vectors 127 G. Virus Strains Nontransmissible by the Vectors 128 III. Discussion 129 A. General Considerations 129 B. Miscellaneous Points 132 C. Some Comparisons of Plant and Animal Tumors 133 References 135 Plant Genetic Tumors H.H. Sirni I. Introduction 138 II. Occurrence of Spontaneous Tumors in Higher Plants 139 A. Within Species 139 B. In Species Hybrids Other than Nicotiana 141 C. In Nicotiana Hybrids 143
7 Contents VII III. Cytogenetic Basis for Tumor Production in Nicotiana Hybrids 144 IV. Morphology and Histology of Nicotiana Tumors 148 V. Effects of Ionizing Radiations and Chemical Agents 149 VI. Physiology, Biochemistry and Tissue Culture 152 A. Chemical Composition of Plants 153 B. Tissue Cultures 154 VII. Synthesis: Genetic Tumors and Gene Regulation 156 References 159 The Relevance of Plant Tumor Systems to an Understanding of the Basic Cellular Mechanisms Underlying Tumorigenesis 165 A. C.Bi λui Subject Index Vols Index Vols
8 Series Editor's Foreword Exaggerated emphasis is placed in today's government research policy planning upon work that deals directly with man and his diseases and with the prevention and treatment of disease in patients. Some of this type of clinical research is undoubtedly good and necessary. However, there remain numerous problems in clinical medicine that no amount of clinical study can solve. For the acquisition of new fundamental knowledge, it will be necessary to resort to experimental and comparative pathology. This includes studies in plants. Let us remember that fundamentally valid laws of genetics were deduced by MENDEL from experiments on plants, and that some of the most widely applicable knowledge in biochemistry today comes from the study of metabolism of yeasts and other microorganisms. It is because of considerations such as these, and with the intention of giving further depth to our search for true progress that we have turned to plant physiologists and pathologists for this volume. F. HoMBURGER, M.D. Series Editor Bío-Research Institute, Inc. Cambridge, Mass. (USA)
9 Guest Editor's Foreword The nonself-limiting neoplastic diseases of plants have had a long and interesting history. As early as 1907, SMιr and TOWNSEND isolated and described a specific bacterium that was found to be responsible for the initiation of a rapidly expanding tumorous disease that was known as the crown gall disease. This discovery attracted considerable interest among pathologists generally because, at the time of that finding, no animal tumor had as yet been produced under defined experimental conditions. The demonstration by ELLERMANN and BANG of the transmissibility of fowl leukemia by means of cell-free filtrates and the experimental production of solid tumors in chickens by Rous were not reported until later. Plant tumors were also among the earliest shown to be transplantable. In 1910, JENSEN who, because of his brilliant investigations on the transplantability of mouse and, later, rat tumors, is commonly considered to be the father of modern experimental cancer research, applied his talents to a study of the transplantability of certain plant tumors as well. As a result of their investigations, both SMιrκ and JENSEN repeatedly expressed the belief that plant and animal tumors have much in common, and JENSEN stated that the plant tumors with which he worked `remind one so much of the malignant tumors of animals, that a closer study of the biological relationships of these tumors would undoubtedly be profitable'. While certain of the less biologically oriented investigators in the field of mammalian oncology may perhaps regard such thoughts as being of questionable value, it should be recognized that the tumor problem is in essence a cellular problem and at that level few differences of fundamental nature exist between members of the two kingdoms. In comparing plant and animal tumors it must be remembered, however, that there are certain structural and functional differences commonly used to distinguish cancers of
10 Guest Editor's Foreword X animals that are more or less restricted to animals and cannot, therefore, be carried over and applied to plants. The most essential characteristic, that of the ability of the cell to grow in an essentially unrestrained or autonomous manner within a host, upon which all other diagnostic features ultimately depend, is, however, equally capable of expression in neoplasia of all higher organisms since it is a characteristic feature of the cell itself. Three nonself-limiting neoplastic diseases of plants which have different and quite distinct proximate causes were selected for discussion here. These diseases were chosen as model systems because they clearly illustrate certain fundamental principles and concepts that may provide insight into the basic cellular mechanisms that underlie neoplastic growth generally. An attempt has been made in the last essay of this volume to relate the significance of the experimental findings in the plant tumor field to an understanding of the tumor problem generally. ARMIN C. BRAUN, Ph. D. Guest Editor The Rockefeller University New York, N.Y. (USA)
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