Advances in Chemotherapy. Volume 2 PDF
Preview Advances in Chemotherapy. Volume 2
Contributors to This Volume E. P. ABRAHAM G. F. GAUSE GERTRUDE B. ELION GEORGE H. HITCHINGS WALLACE FOX G. G. F. NEWTON EMIL FREI, III J. E. PEACHEY EMIL J. FREIREICH F. C. PEACOCK Advances in Chemotherapy Edited by ABRAHAM GOLDIN F. HAWKING National Cancer Institute National Institute for Medical Research National Institutes of Health Mill Hillf London U.S. Public Health Service England Bethesda, Maryland ROBERT J. SCHNITZER Formerly, Chemotherapy Department Hoffmann—LaRoche Inc. Nutley, New Jersey VOLUME 2 1965 Academic Press · New York and London COPYRIGHT © 1965, BY ACADEMIC PRESS INC. ALL RIGHTS RESERVED. NO PART OF THIS BOOK MAY BE REPRODUCED IN ANY FORM, BY PHOTOSTAT, MICROFILM, OR ANY OTHER MEANS, WITHOUT WRITTEN PERMISSION FROM THE PUBLISHERS., ACADEMIC PRESS INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS INC. (LONDON) LTD. Berkeley Square House, London W.l LIBRARY OF CONGRESS CATALOG CARD NUMBER: 64-21671 PRINTED IN THE UNITED STATES OF AMERICA List of Contributors Numbers in parentheses indicate the pages on which the authors' contributions begin. E. P. ABRAHAM, Sir William Dunn School of Pathology, University of Oxford, England (23) GERTRUDE B. ELION, Wellcome Research Laboratories, Burroughs Wellcome and Co., U.S.A., Inc., Tuckahoe, New York (91) WALLACE FOX, Tuberculosis Research Unit, Medical Research Coun- cil, Holly Hill, Hampstead, London, England (197) EMIL FREI, III, * Medicine Branch, National Cancer Institute, National Institutes of Health, Bethesda, Maryland (269) EMIL J. FREIREICH, ^Medicine Branch, National Cancer Institute, National Institutes of Health, Bethesda, Maryland (269) G. F. GAUSE, Institute of New Antibiotics, Moscow, U.S.S.R. (179) GEORGE H. HITCHINGS, Wellcome Research Laboratories, Bur- roughs Wellcome and Co., U.S.A., Inc., Tuckahoe, New York (91) G. G. F. NEWTON, Sir William Dunn School of Pathology, University of Oxford, England (23) J. E. PEACHEY, f Rothamsted Experimental Station, Harpenden, Eng- land (1) F. C. PEACOCK, Jealott/s Hill Research Station, Bracknell, England (1) * Present address: University of Texas M. D. Anderson Hospital and Tumor Institute, Houston, Texas, f Present address: Commonwealth Bureau of Helminthology, The White House, St. Albans, England. ν Preface The general aims of Advances in Chemotherapy were stated in the Preface of Volume 1. As was indicated therein, this serial publication is designed to provide comprehensive and authoritative surveys of progress in all fields of experimental and applied chemotherapy. In this volume Peacock and Peachey discuss the chemical control of nematodes in plants; this article will draw the attention of chemists and medical biolo- gists to a field in which conditions are sometimes similar and sometimes very different from those in man and animal. Abraham and Newton give a detailed account of the antibiotics of the cephalosporin group—their chemistry, biochemistry, and anti-infectious properties. The antibiotics described in Gause's article are members of the olivomycin group; they are of interest because of their antineoplastic activity. Chemotherapeutic action against malignancies is an important part of the contribution by Elion and Hitchings on the metabolic basis of the action of purines and pyrimidines. The specific relation of these structures to nucleic acids and cell nuclei extends the range of activity of these compounds to viral and protozoan infections. The application of chemotherapy to neo- plastic diseases of man is described in the article by Frei and Freireich who analyze the complex field of drug response in acute leukemia. The special problems which are involved in the chemotherapy of tuberculosis as applied in developing countries, particularly in India, are discussed by Fox. It is hoped that these reviews will be useful to all concerned in pro- viding convenient summaries of information scattered throughout many different publications and in stimulating the interest of workers in other disciplines. The Editors are indebted to the contributors for their willingness to publish the results of their wide experience in the various fields of chemo- therapy. The expert cooperation of the members of the staff of Academic Press is gratefully acknowledged. A. GOLDIN F. HAWKING October, 1965 R. J. SCHNITZER vii Systemic Control of Plant Nematodes F. C. PEACOCK AND J. E. PEACHEY Jealotfs Hill Research Station, Bracknell, England, and Rothamsted Experimental Station, Harpenden, England I. INTRODUCTION 1 II. CHEMICALS MOBILE IN PLANTS 2 A. Inorganic Salts 3 B. Sugars 3 C. Nitrogenous Compounds 3 D. Growth Regulators 4 III. BIOCHEMISTRY OF NEMATODE DISEASE 5 A. Respiration 5 B. Nutrition of Host and Parasite 5 C. Antimetabolites and Enzymes 6 D. Auxins 8 IV. NATURALLY ARISING RESISTANCE FACTORS 9 V. CHEMOTHERAPEUTIC TREATMENTS 11 A. Chemical Treatment of Dormant Plant Material ... 11 B. Systemic Treatment of Growing Plants 13 VI. CONCLUSIONS 18 REFERENCES 19 I. INTRODUCTION Nematode diseases of plants are at present chemically controlled in practice largely by chemical sterilization or surface disinfection of the soil (Goring, 1962; Peachey, 1963), carried out as a preventive treat- ment before planting. The purpose of this review is to discuss the aims 1 2 F. C. PEACOCK AND J. Ε. PEACHEY of, and progress made toward, eradicant or protective chemical treat- ment of the plant. An effective chemical may protect the plant by inactivating attractive root exudates, or by killing or repelling nematodes before they invade; or it may be truly therapeutic, killing nematodes in the plant by direct toxicity or by inducing a resistant reaction in the plant. Toxicity to nematodes may not be an essential property; but a chemotherapeutant must be able to enter and move systemically within the plant, without harming it, and must alter the host/parasite relationship in favor of the plant. In considering the progress already made and the chances of finding new and more effective chemicals of this nature, it is necessary to draw on what is known of (i) translocating systems in the healthy plant, (ii) biochemical explanations of disease symptoms, (iii) the nature of re- sistance in plants to disease, and (iv) the mobility of known toxicants introduced artificially into the plant. II. CHEMICALS MOBILE IN PLANTS Substances may move in plants either in the phloem or in the xylem. In general, chemicals which move in the xylem are carried on the transpiration stream and move up from the roots to the shoots; this process is independent of living cells and is governed by the solubility of the chemical in soil water, by adsorption on cell walls, and by the rate of transpiration of the plant. Plant nutrients and many soil-applied toxicants move in this way. Selective accumulation in different parts of the plant depends upon physical properties of the compound and may be made use of in chemotherapeutic treatment; lipoid-soluble chemicals tended to accumulate in roots (Crowdy and Rudd Jones, 1956) and a positive charge assisted retention (Edgington and Dimond, 1960). Chemicals not retained in roots tended to move most rapidly to points of maximum transpiration, such as young but fully expanded leaves (Biddulph et al., 1958). Movement in the phloem depends upon the activity of living cells, and the type of chemical moved is highly specific. Accumulated photosynthates move out of mature leaves to regions of rapid growth, such as developing leaves and fruit, roots and tubers. Lateral movement also may occur between xylem and phloem, and dif- fusion into surrounding parenchyma has been recorded (Tietz, 1954). Under certain environmental conditions movement from shoot to root Systemic Control of Plant Nematodes 3 may occur in xylem (Clor et al., 1963). Current thought on circulatory systems in plants has been reviewed by Bollard (1960), Zimmermann (1960, 1961), Kursanov (1963), and Nelson (1963). Certain chemicals sprayed onto leaves may enter and be distributed within the plant to a greater or lesser extent—fertilizers and trace ele- ments in solution are commonly applied in this way. Not enough is known of the chemical and physical properties governing leaf penetra- tion and mobility in the plant to be able to predict systemic activity of this kind. However, four main groups of compounds are known to move in phloem; these are inorganic salts, sugars, nitrogenous compounds, and growth regulators. A. Inorganic Salts Nutrient elements have been classified according to their degree of mobility in bean plants from leaf application. Bukovac and Wittwer (1957) regarded Rb, K, Na, P, CI, and S as mobile; Zn, Cu, Mn, Fe, and Mo as partially mobile; and Ca, Sr, and Ba as immobile. Radio- activity was detected in Heterodera rostochiensis and Meloidogyne in- cognita in the roots of tomato plants fed a nutrient solution containing P32-labeled phosphoric acid (Dropkin and King, 1956), and in Longi- dorus maximus feeding on the roots of sugar beet whose leaves had been sprayed with the monosodium salt of the labeled acid (Sprau and Suss, 1962). B. Sugars Many products of photosynthesis accumulate in leaves and vary widely according to plant species and environment. The principal substance moving out of leaves via the phloem is sucrose, which can account for up to 90% of all metabolic products being transported. Smaller quantities of related nonreducing sugars also occur, but may not move in this form (see Swanson, 1959; Kursanov, 1963). Sucrose applied to leaves has been shown to aid translocation of other compounds (Barrier and Loomis, 1957) and this mechanism is said to be enhanced by boron (Gauch and Dugger, 1953). C. Nitrogenous Compounds Reinbothe and Mothes (1962) regarded allantoic acid and allantoin as the forms in which most naturally occurring nitrogenous matter is 4 F. C. PEACOCK AND J. Ε. PEACHEY transported in the phloem, and in which it occurs in root exudates. At different times of year and in different plant species, varying numbers and amounts of as many as 12 amino acids may occur, particularly glu- tamic, aspartic, and α-aminobutyric acids, together with the amides of serine, leucine, valine, proline, and alanine (Kursanov, 1963). Young roots of corn and sunflower immersed in water excreted large amounts of amino acids and amides, sugars, and organic acids (Grineva, 1962). When amino acids were applied to the leaves of soybean, tomato, and wheat growing in a dilute mineral solution, measurably increased amounts of the same amino acids appeared in the growth medium ( Katz- nelson et al., 1954). D. Growth Regulators Little is known of the transport in plants of naturally occurring growth- regulating compounds, but considerable work has been done with synthetic compounds. The observation by Preston et al. (1954) that α-methoxyphenylacetic acid (I), applied to leaves of one bean plant, was transported to the roots, exuded, and taken up by the roots of an adjacent plant suggested a new approach to systemic control of root parasites. Mandelic acid moved as readily but had little growth-modi- fying effect (Mitchell, 1955). Subsequently a further 29 compounds closely related to α-methoxyphenylacetic acid were reported to have similar biological properties (Mitchell et al., 1959). Clor and Crafts (1957) compared the movement of 2,4-dichlorophenoxyacetic acid (2,4- OCHCOOH 2 (IV) (V) (VI) Systemic Control of Plant Nematodes 5 D) (II), aminotriazole (III), and urea out of treated leaves; all three were transported to the roots; 2,4-D was exuded from the roots, whereas aminotriazole and urea tended to be retained there. Maleic hydrazide (IV) was more mobile than 2,4-D, aminotriazole, urea, or monuron (3-p-chlorophenyl-l,l-dimethylurea) (V), according to Crafts and Yama- guchi (1958). Trichloro- and tetrachlorobenzoic acids also moved from the leaves of treated bean plants into adjacent untreated plants via the root systems (Linder et al., 1958). Certain growth regulators are also of interest for their effect on the translocation of other materials; thus transport of P32 was increased by giberellic acid (Linck and Sudia, 1960). Naturally arising indole-3-acetic acid (VI) and artificially applied 2,4-D enhanced the transport of or- ganic substances in the phloem (Kursanov, 1963). ΠΙ. BIOCHEMISTRY OF NEMATODE DISEASE Krusberg (1963) has recently reviewed present knowledge of plant responses to nematode infection. By comparison with the extensive studies of fungal, viral, and bacterial disease presented by Horsfall and Dimond (1959-1960), relatively little quantitative work has been done. Gross symptoms of nematode disease—browning of damaged tissue, wilt- ing and yellowing of leaves, and galling of roots—are similar to those of nonparasitic infection, and similar metabolic changes might be expected. A. Respiration The respiration rate of diseased tissue is usually greater than that of normal tissue. Krusberg (1963) quotes several instances (Turlygina, 1957; Nonaka, 1959) of increased respiration in nematode disease. Bird and Millerd (1962) were unable to show increased respiration in to- mato roots galled by M. incognita, and suggested that hypertrophied cortical cells may respire at a lower than normal rate. B. Nutrition of Host and Parasite Infection of a susceptible host by nematodes often leads to a local stimulation of growth of plant cells surrounding the parasite, and an increased flow of materials into these cells. At the same time nutrients
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