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Handbuch der experimentellen Pharmakologie Vo 1. 48 Heffter-Heubner New Series
Handbook of Experimental Pharmacology
Herausgeber Editorial Board G. V R. Born, Cambridge· O. Eichler, Heidelberg A. Farah, Rensselaer, NY· H. Herken, Berlin A. D. Welch, Memphis, TN
Beirat AdvisOlY Board E.J.Ariens· Z.M.Bacq· P.Calabresi· S.Ebashi· E.G. Erdos V. Erspamer . U. S. von Euler· W. S. Feldberg· G. B. Koelle· O. Krayer T.A. Loomis· H. Raskova . M. Rocha e Silva· F. Sakai· J. R.Vane P. G.Waser . W. Wilbrandt
Arthropod Venoms Contributors
D.WAlsop . R. L. Beard· S. Bettini· M. S. Blum P. M. Brignoli . W Bucherl . A. Delgado Quiroz· C. R. Diniz H. Edery . P. Efrati . T. Eisner· S. Gitter· M. Goyffon M. R. Gray . Y. Hashimoto . H. R. Hermann· K. Hicks J. Ishay . H. Joshua· S. Konosu . J. Kovoor . G. Levy Z. Mareti6 . M. Maroli . J. Meinwald . A. Minelli F. Miranda· M. F. Murnaghan· R. O'Connor· F. J. O'Rourke M. L. Peck· J. E. Percy· F.A. Pereira-Lima· W Rathmayer H. Rochat . L. M. Roth . S. Schenberg . H. Schenone A. Shulov . H. L. Stahnke· G. Suarez· S. K. Sutherland J. Weathers ton . E. Zlotkin
Editor
Sergio Bettini
Springer-Verlag Berlin Heidelberg New York 1978
Prof. Dr. S. BETTINI, Istituto Superiore di Sanita, Viale Regina Elena 299, Rome/ Italy
ISBN-13: 978-3-642-45503-2 DOl: 10.1007/978-3-642-45501-8
e-ISBN-13: 978-3-642-45501-8
Library of Congress Cataloging in Publication Data. Main entry under title: Arthropod venoms. (Handbook of experimental pharmacology: New series: v. 48) Bibliography: p. Includes index. I. Arthropoda, Poisonous-Venom. J. Alsop. David W .. 1939- II. Bettini, Sergio, 1916- III. Series: Handbuch der experimentellen Pharmakologie: New series: v. 48. QP905.H3 vol. 48 [QP94I.A73]615'.1'08s [595'.2'0457]77-24628
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© by Springer-Verlag Berlin/Heidelberg 1978.
Softcover reprint of the hardcover 1st edition 1978
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
Arthropod venoms have received much attention and have played an important role in folklore and medicine since ancient times. Scorpion envenomation, "tarantism," bee and wasp stings are among those subjects about which most has been speculated and written in the past. In the last 50 years or so, a great number of scientific papers have been devoted to arthropod venoms, but only a few volumes have been designed to collect this rapidly increasing material, and these are not recent. Of late, the chemistry and mode of action of several arthropod venoms have been thoroughly studied, and some of these substances will probably be used as pharmacological tools and also as therapeutic agents.
The aim of the present volume is to collect in manual form new information as well as the old notions on arthropod venoms.
Even though it was our intention to present a volume on arthropod venoms, and not on venomous arthropods, inevitably we were forced to include information on venom-producing organisms as well. We assumed, in fact, that those scientists for whom the present manual is primarily intended (biochemists, particularly comparative biochemists, and pharmacologists) should be familiar with the biologic elements concerning the venom-producing species; which should show them how important it is to operate in close collaboration with biologists specialized in venomous arthropod systematics and biology. Furthermore, the distribution, ethology, and ecology of venomous species should be thoroughly known if the specific therapy and prevention of envenomation are to be correctly undertaken by clinicians and public health specialists respectively. For these reasons the reader, besides an introductory chapter on arthropod systematics, will find in the first part of each chapter much space devoted to the distribution and biology of the venomous species.
The origin and nature of animal substances called venoms and the concept of toxicity have been much debated throughout the years and several definitions of the term "venom" have been given by different authors. In our volume the proposed definition is the old one of E.N. PAWLOWSKY (Gifttiere und ihre Giftigkeit. Jena: G. Fischer, 1927), later adopted by E. KAISER and E. MICHL (Die Biochemie der tierischen Gifte. Wien: F. Deuticke, 1958) which reads as follows: "Those substances are considered venoms which, due to their chemical characteristics, after their penetration into the human or animal body can cause, even in small doses, alterations in their health, or death." By following such a definition, the authors were allowed to include in this volume topics regarding all toxic substances derived from arthropods, even those not originating from venomous apparatuses.
The concept of toxic properties of tissues has been well treated already by M. PHISALIX (Animaux Venimeux et Venins. Paris: Masson, 1922) and recently
VI Preface
by E. HABERMAN (Chemistry, Pharmacology, and Toxicology of Bee, Wasp, and Hornet Venoms. In: Animals and their Venoms, Vol. III, Eds. W. Bucher! and E. Buckley. New York: Academic Press, 1971). Obviously the limits of this concept are vague: many substances present in arthropods, for instance, may cause toxic phenomena if administered in high doses. As HABERMAN states, the frontier between "simple" body constituents and "venom" is not sharply marked: an example is given by biogenic amines which are present both in tissues and in venoms. On the other hand, in some cases (e.g., bee venom) when the venom is administered at very low doses, the original sense of the term gives way to its conceptual opposite, that of medicament.
We have deemed it useful to broaden the subject by also including in the volume the arthropod defensive substances, some of which are not to be considered as toxic sensu stricto, even though they do induce profound behavioral modifications in other species. This type of compound appears more puzzling if, for instance, we consider that some ketones are utilized by ants both for defensive and communicative functions (M.S. BLUM and H.R. HERMAN, Chapter 25).
The limits to be attributed to the term venom could not, therefore, be fixed a priori, but were left in each case to the judgment of the individual authors.
As initially planned, the volume should have also included chapters dealing with species belonging to orders less known for their poisonous characteristics. It is well known that the saliva of Diptera, for instance, often causes local and general toxic phenomena not to be attributed to allergic responses only (e.g., toxic reactions due to bites of tabanid larvae, asilids, simulids, ceratopogonids, etc.). Information in such a field, however, is so scanty that we had to abandon the idea. On the other hand, a great deal has already been published on insect allergy, a topic more pertinent to the field of immunology. On this subject, excellent reviews have been presented by S. SHULMAN (Allergic responses to insect bites. Annual Rev. Entomol. 12,323-346,1967), B.F. FEINGOLD, E. BENJAMINI and D. MICHAELI (The allergic responses to insect bites. Annual Rev. Entomol. 13, 137-158, 1968) and c.A. FRAZIER (Insect Allergy, St. Louis: W.H. Green, 1969), to which the reader is referred.
The volume has been divided into 26 chapters, each one related to a specific zoologic group, this being the only possible way of arranging the abundant material on the various subjects without risking many omissions or repetitions. Unfortunately, as will be noted by the careful reader, a few were unavoidable. The existence of a very large number of arthropod-borne toxic substances, and of even larger number of specialists, 43 of whom were invited to participate in the preparation of the present volume, made this drawback inevitable.
In order to cover the naturalistic as well as the biochemical and pharmacologic aspects of each chapter, more than one author was often needed. Though this represented an advantage as far as the thorough treatment of the subject was concerned, it negatively influenced the uniformity of the volume because of the diverse nature and amount of information available for each topic.
The authors of the volume are among the most qualified specialists in their fields. They were asked to include in their chapters all that is known on the subject, so as to match the purpose and tradition of the monographs published by the Handbook of Experimental Pharmacology.
Preface VII
Editing of the present volume started in January 1974. The mailing of the first group of invitations coincided with one of the most serious postal strikes in Italy, so that a great deal of mail went astray. This, plus the courteous declining by some scientists and the dehiy of others in providing the manuscripts caused the volume to be published at the end of 1977 and consequently a few of the chapters may be somewhat outdated.
I wish to express my gratitude to all authors who have taken on the burden of this task with enthusiasm and skill: they should be praised for the high standard of their reviews. Thanks are due to Professor P. BRIGNOLI for the time he has devoted to discussing some of the naturalistic problems of the volume. I am particularly indebted to Miss A.M. LOPOMO for her excellent secretarial work. I also wish to thank member of the staff of Springer-Verlag, who were extremely cooperative in the preparation of the volume.
In January of this year we received the very sad news that our Japanese colleague Professor Y. HASHIMOTO, who had prepared jointly with Professor S. KONOSU the chapter "Venoms of Crustacea and Merostomata", had died on 23 September 1976. I am sure that I share the feelings of all the other authors in expressing our deep sorrow and sympathy to Professor HASHIMOTO'S family and to all those who have had the pleasure of working with him.
August 1977 SERGIO BETTINI
Table of Contents
CHAPTER 1
Introduction to Venomous Arthropod Systematics. P.M. BRIGNOLI
A. Introduction . . . . . . . . . . . . . I. What are Arthropoda? . . . . .
II. The Main Divisions of the "Type" . III. The Chelicerata . . . . . .
1. Scorpionida . . . . . . 2. Uropygi or Thelyphonida 3. Pseudo scorpionida . 4. Opiliones. 5. Acarina .. 6. Araneae ..
IV. The Crustacea. V. The" Myriapoda" .
1. Chilopoda . . . 2. Diplopoda . . .
VI. The Hexapoda or Insecta. 1. Blattodea and Dermaptera 2. Rhynchota and Anoplura 3. Aphaniptera . 4. Coleoptera . . 5. Hymenoptera . 6. Diptera ... 7. Lepidoptera .
VII. Some General Advices 1. How to Identify an Arthropod 2. How to Conserve an Arthropod. 3. What to Expect from the Bibliography .
References
CHAPTER 2
Venoms of Crustacea and Merostomata. Y. HASHIMOTO and S. KONOSU. With 10 Figures
A. Introduction . . . . . . . . . . . . . . B. Crustaceans Suspected of Being Poisonous .
1 1 1 2 2 2 3 3 3 3 4 4 4 4 5 5 6 6 6 6 6 6 7 7 7 7
8
13 14
x
C. Toxicity of Crabs. . . . . . . . . . I. Crabs Containing Saxitoxin. . .
II. Toxicity of Lophozozymus pic tor .
D. Biology of Poisonous Xanthid Crabs I. Zosimus aeneus
1. Description. . . . . . . 2. Color in Life . . . . . . 3. Habitat and Distribution. 4. Feeding Habits and Spawning Season
II. Platypodia granulosa 1. Description. . . . . . . 2. Color in Life . . . . . . 3. Habitat and Distribution.
III. Atergatis floridus. 1. Description. . . . . . . 2. Color in Life . . . . . . 3. Habitat and Distribution.
IV. Lophozozymus pic tor 1. Description. 2. Coloration . . . 3. Distribution . .
E. Chemistry of Toxins in Crabs I. Purification of Z. aeneus Toxin
II. Chemical Properties of Z. aeneus Toxin. III. Chemical Properties of Toxins from Other Crabs.
F. Pharmacology of Toxins in Crabs .... I. Z. aeneus Toxin . . . . . . . . .
1. Poisoning Cases and Symptoms . 2. Signs in Test Animals . . . 3. Effects on Nerve Excitation.
II. The Toxin of L. pictor III. Toxin in Other Crabs. . .
G. Coconut Crab Poisoning. . . . I. Biology of Coconut Crab .
1. Description. . . . . . 2. Color when Alive, Habitat, and Distribution
II. Toxicity of the Coconut Crab
H. Horseshoe-Crab Poisoning. . . . I. Biology of Horseshoe Crabs.
II. Toxicity of Horseshoe Crabs III. Clinical Characteristics . . .
I. Prevention and Treatment of Crab Poisoning.
References . . . . . . . . . . . . . . . . .
Table of Contents
15 15 19
19 19 20 20 20 20 20 21 22 22 22 22 22 24 24 24 24 24
24 25 26 26
27 27 27 27 28 32 33,
33 34 34 34 34
35 35 36 37
37
37
Table of Contents XI
CHAPTER 3
Defensive Secretions of Millipeds. T. EISNER, D. ALSOP, K. HICKS, and J. MEINWALD. With l3 Figures
Introduction. . . . . . . . . . . . . . . . . . . . . . . . A. Distribution, Structure, and Mode of Operation of the Glands B. Chemistry of the Secretions . . . . C. Biochemistry of the Secretions . . . D. Defensive Effectiveness of the Glands E. Implications to Humans. . F. Other Defenses of Millipeds
References . . . . . . . . .
CHAPTER 4
Secretions of Centipedes. A. MINELLI. With 2 Figures
A. The Centipedes. . . . . . . . . . . . . . . . B. Glands and Secretions. . . . . . . . . . . . .
I. Defensive and Offensive Weapons of Centipedes. II. Topography of the Exocrine Glands
III. Histology of the Glands . . . . . . . . . . . 1. The Forcipular Gland .......... . 2. Ventral and Coxal Glands of the Geophilomorphs .
IV. Secretions. . . . . . . . 1. Sticky Secretions . . . 2. Luminescent Secretions 3. Smelling Secretions . . 4. Venom of the Forcipular Glands
a) Effects of Bite on Humans. . b) Effects of Bite on Laboratory Animals. c) Chemistry of the Venom.
References
CHAPTER 5
41 42 51 55 58 65 66
67
73 75 75 75 76 76 77 78 78 79 79 80 80 81 82
82
Secretions of Opilionids, Whip Scorpions and Pseudoscorpions. T. EISNER, D. ALSOP, and J. MEINW ALD. With 7 Figures
A. Order Opiliones . . . B. Order U ropygi . . . . C. Order Pseudoscorpiones
References . . . . . . .
87 94 97
97
XII Table of Contents
CHAPTER 6
Review of the Spider Families, with Notes on the Lesser-Known Poisonous Forms. S. BETTINI and P.M. BRIGNOLI
A. Orthognatha = Theraphosomorphae = Mygalomorphae . B. Labidognatha = Araneomorphae
I. Haplogynae. . . . . . . II. Entelegynae = Trionychae .
I. Araneoidea. . . . . . a) Araneidae = Argiopidae b) Linyphiidae c) Theridiidae
2. Lycosoidea. . a) Agelenidae. b) Argyronetidae c) Pisa uridae . d) Lycosidae . . e) Oxyopidae. .
III. Entelegynae=Dionychae I. Gnaphosidae (= Drassodidae = Drassidae) 2. Clubionidae . . . . . . . . . 3. Ctenidae. . . . . . . . . . . 4. Eusparassidae (= Heteropodidae) 5. Thomisidae. 6. Salticidae.
IV. Cribellatae
References . . . . .
CHAPTER 7
Venoms of Dipluridae. M.R. GRAY and S.K. SUTHERLAND
Introduction. . . . . . . . . . . . . . . . . . . .
A. Systematics and Distribution. M.R. GRAY. With 1 Figure.
B. Biology and Venoms. S.K. SUTHERLAND. With 10 Figures.
I. Introduction II. Notes on Atrax robustus . . . . . . . . .
III. Source of Venom for Experimentation ... 1. Maintenance of Spiders in the Laboratory 2. Collection of Venom and Venom Yields .
IV. Structure of the Venom Glands in Atrax robustus V. Toxicity of Venom in Various Animal Species.
1. Atrax robustus Venom ..... 2. Venom of Other Species of Atrax .....
103 108 108 110 110 110 III III III 112 112 112 112 114 114 114 114 116 116 116 117 117
118
121
121
126
126 127 129 129 130 130 132 132
" 133
Table of Contents XIII
VI. Chemistry of Sydney Funnel-Web Spider (Atrax robustus) Venom 134 VII. Pharmacopathologic Studies with Atrax robustus Venom 137
1. Intact Animal Experiments. . . . 137 2. Studies Upon Isolated Preparations 139 3. Investigation of Antidotes . . . . 141
VIII. Immunologic Studies on Atrax Venom 142 IX. Human Envenomation by Atrax robustus 143
1. General . . . . . . . . . . . . . 143 2. Signs and Symptoms of Envenomation. 144 3. Treatment of Victims 145 4. Prognosis 146
XD~m~oo 1%
References . 147
CHAPTER 8
Venoms of Theridiidae, Genus Latrodectus
A. Systematics, Distribution and Biology of Species; Chemistry, Pharmacology and Mode of Action of Venom. S. BETTINI and M. MAROLI. With 8 Figures
I. Systematics and Distribution of Species. 149 II. Biology. . 151
1. Habitat . 151 2. Habits. . 152 3. Life Cycle 152 4. Biting Act 154
III. Venomous Apparatus 154 IV. Venom. . . . . . . 156
1. Methods for Obtaining the Venom 156 2. Chemistry . . . . . . . . . . . 157 3. Separation of Venom Components. 158
V. Toxicity of Venom in Different Animal Species 159 1. Whole Venom . . . . . . . . . . . . . 159 2. Venom Components. . . . . . . . . . . 160
VI. Pharmacology and Mode of Action on Different Zoological Groups 162 1. Invertebrates 166
a) Crustacea 166 b) Insecta . 167 c) Mollusca 170
2. Vertebrates. 170 a) Pisces. . 170 b) Amphibia 170 c) Aves . . 172 d) Mammalia . 172
3. Toxicity in Cells Cultivated in vitro 177 4. Effect of Toxin on Lipid Bilayer Membranes 177
XIV Table of Contents
5. Distribution of Venom in Envenomed Animals . . 177 6. Protective Activity of Venom Against Botulin Toxin. 178
VII. Toxicity of Eggs and Tissue Extracts 178
References . . . . . . . . . . . . . . . . . . . . . . . . 179
B. Epidemiology of Envenomation, Symptomatology, Pathology and Treatment. Z. MARETIC. With 3 Figures
I. Epidemiology . . . . . . . 185 1. Geographical Distribution 185
a) Europe 185 ~A~ 1~ c) Africa. . 186 d) America . 186 e) Australia 187
2. Factors Influencing Number of Spiders and Frequency of Contacts with Man. Epidemics of Latrodectism 187
3. Other Epidemiological Data 191 II. Human Symptomatology. . . . . . . 192
III. Differential Diagnosis . . . . . . . . 199 IV. Clinical Course, Duration of Hospitalization, Convalescence, and
Complications. . 200 1. Course. . . . 200 2. Convalescence 200 3. Complications 201
V. Prognosis. . . . 201 VI. Post-Mortem Findings 201
VII. Aspecific Therapy . . 202 VIII. Specific Therapy. . . 204
1. Preparation of Sera 204 2. Cross Activity of Sera Obtained with Antigens from Different
Species of Spiders. . . . . . . . . . . 206 3. Time and Dose of Serum Administration. 206
IX, Prevention of Latrodectism 207
References . 207
CHAPTER 9
Venoms of Theridiidae, Genus Steatoda. Z. MARETIC. With 2 Figures
A. Introduction . . . . . . . . . . . . . 213 B. Venomous Spiders of the Genus Steatoda . . . . . . . . . . 213
I. Steatoda paykulliana . . . . . . . . . . . . . . . . . 213 1. Distribution, Biology, Morphology, and Venomous Apparatus 213 2. Effects of the Toxin 214
II. Steatoda grossa 216
References . . . . . . . . . 216
Table of Contents
CHAPTER 10
Venoms of Ctenidae. S. SCHENBERG and F.A. PEREIRA LIMA. With 15 Figures
A. Introduction . B. Morphology of the Venomous Apparatus
I. Chelicerae 1. Fangs or Claws . 2. Basal Segment 3. Musculature
II. Venom Glands 1. M uscula ture 2. Histology of the Glandular Tissue .
C. Methods of Collecting Venoms. 1. Method of Collecting Venom by its Extraction from the Glands. 2. Method of Collecting Venom by Electrical Stimulation.
D. Physiology of Biting. E. Crude Venom Pharmacology .
I. Effect on Dogs 1. Excruciating Local Pain 2. Sneezing . 3. Lacrimation and Mydriasis. 4. Hypersalivation . 5. Erection . 6. Ejaculation. 7. Toxicity 8. Hypotensive Response 9. Tachyphylaxis
II. Guinea Pig Ileum Contraction. III. Effects on Mice
1. Local Pain. 2. Hypersalivation . 3. Erection . 4. Ejaculation . 5. Toxicity 6. Distensive Paralysis
IV. Effects on Other Animals .
F. Biochemistry of the Phoneutria nigriventer Venom, and its Fractionation . . . . . . . . . . .
Procedures for
I. Immunologic Aspects of the Venom . . ..... II. Enzymic Inactivation. . . . . . . . . . . . . .
III. Physicochemical Properties of the Active Polypeptides IV. Dialysis. . . . . . . . . . . . . . . . . . . . . V. Venom Fractionation. . . . . . . . . . . . . . .
1. Ammonium Sulfate, Electrophoresis, Barium Sulfate Adsorption 2. Flaccid Paralysis . 3. Hemorrhagic Effect . . . . . . . . . . . . . . . . . . . .
xv
217 219 219 219 220 220 221 221 221
222 223 223
224 224 225 225 226 226 226 226 228 229 228 229 229 229 230 230 231 231 231 232 232
233 233 233 234 235 236 236 236 237
XVI
VI. Gel Chromatography. . VII. Ion Exchange Columns.
Table of Contents
237 240
G. Envenomation in Man. . 242 242 243 243
I. Symptomatology. . . . II. Frequency of Cases . .
III. Therapy of Envenomation
References . . . . . . . . . . . 244
CHAPTER 11
Venoms of Scytodidae. Genus Loxosceles. H. SCHENONE and G. SUAREZ. With 6 Figures
A. Distribution and Biology of Venomous Species. Chemistry, Toxicity, Phar-macology and Mode of Action of Venom . . 247
I. Systematics and Distribution of Species. . . . . . . 247 II. Morphology and Biology . . . . . . . . . . . . . 248
III. Anatomy and Histology of the Venomous Apparatus. 249 IV. Physiology, in Particular Physiology of the Bite 250 V. Chemistry of the Venom . . . . . . . . . 251
l. Methods of Obtaining the Venom. . . . . 251 2. Chemical Composition of Venom Extracts . 252 3. Fractionation and Characterization of Venom Proteins. 252 4. Enzymatic Properties of Venom. . . . . . . . . . . 253 5. Immunogenicity of the Venom . . . . . . . . . . . 254
VI. Toxicity of Venom and other Components of Loxosceles on Various Animal Species . 254 1. In vivo Studies . . . . . . . . . . 254 2. In vitro Studies . . . . . . . . . . 254
VII. Pharmacology and Mechanism of Action 260
B. Epidemiology, Symptomatology, Pathology, Prognosis, Treatment, and Prevention of Envenomations . . . . . . . . . . . . . . . . . . . 260
I. Epidemiology of Envenomations. Geographic and Temporal Distri-bution of Cases. Other Epidemiologic Data 260
II. Human Symptomatology . . . . . . . . 262 l. Cutaneous Form of Loxoscelism . . . 262 2. Viscerocutaneous Form of Loxoscelism 264
III. Differential Diagnosis . . 266 l. Cutaneous Forms. . . . . . . . . . 266 2. Viscerocutaneous Form . . . . . . . 266
IV. Clinical Course and Duration of Hospitalization. 266 l. Local Necrosis and Ulcer. . . . . . . 266 2. Viscerocutaneous Involvement . . . . 266
V. Prognosis and Frequency of Lethal Cases . 266 VI. Pathology. . . . . 267
VII. Laboratory Findings . . . . . . . . . . 268
Table of Contents
VIII. Treatment 1. Nonspecific. 2. Specific
IX. Prevention
References . . . .
CHAPTER 12
XVII
269 269 270 271
271
The Genus Centruroides (Buthidae) and Its Venom. H.L. STAHNKE. With 2 Figures
A. Species and Their Distribution . . . . . B. Some Aspects of Behavior . . . . . . . . . . . . C. The Prevention and Control of Scorpions . . . . . D. Morphology and Function of the Venom Apparatus E. Extracting Venom . . . . . . . . . . . . . F. Mammalian Physiologic Reaction to the Venom . .
I. Clinical Observations. . . . . . . . . . . . 1. Symptoms of Severe Envenomization (Children under 16
of Age) ............. . 2. Symptoms of Envenomization in Adults
II. Preclinical Observations . . . . 1. Pharmacologic Considerations 2. Pathologic Effects. . . 3. Allergic Potentialities .
G. Human Behavior and Scorpions H. Venom Chemistry. . . . 1. Suggestions for Therapy.
References . . . . . . . .
CHAPTER 13
Venoms of Buthinae
A. Systematics and Biology of Buthinae. A. SHULOV and G. LEVY. References . . . . . . . . . . . . . . . . . . . . . . .
B. Symptomatology and Treatment of Buthinae Stings. P. EFRA TI
I. Pathophysiology of the Envenomation II. Treatment
References . . . .
279 280 282 284 286 287 287
Years 287 287 287 287 288 289
289 289 294
294
309 312
312
314 316
316
XVIII Table of Contents
C. Chemistry and Pharmacology of Buthinae Scorpion Venoms. E. ZLOTKIN, F. MIRANDA and H. ROCHAT. With 11 Figures
I. Introduction . . . . . . . II. Chemistry. . . . . . . . . .
1. Isolation and Purification 2. Composition and Structure . 3. Chemical Modifications . .
III. Pharmacology. . . . . . . . 1. Lethal Potency and Symptomatology 2. Cardiovascular Effects. . .
a) Clinical Investigations. . b) Histopathologic Changes c) Vascular Effects . d) Cardiac Dynamics
3. Respiratory Effects . 4. Muscular Effects . .
a) Direct Effects on Skeletal Muscles b) Effects Through Neuromuscular Junctions. c) Effects on Smooth Muscles
5. Action on Axonal Membranes 6. Action on Arthropods . . . .
IV. Action Mechanisms . . . . . . 1. Autopharmacologic Intoxication 2. Is There a Direct Excitatory Effect on Muscles? . 3. Interactions with Sodium, Calcium, and TTX .. 4. The Pharmacologic Diversity of Scorpion Toxins
References
CHAPTER 14
Venoms of Tityinae
317 318 319 323 328 328 328 330 330 330 332 334 336 337 338 338 340 341 345 351 351 353 354 356
362
A. Systematics, Distribution, Biology, Venomous Apparatus, etc. of Tityinae; Venom Collection, Toxicity, Human Accidents and Treatment of Stings. W. BDCHERL.
With 3 Figures
1. The Venomous Species and Their Geographical Distribution 371 2. Description of the Most Venomous Species. 374 3. Frequency of the Most Venomous Species 375 4. Food and Life Habits . . . . . . . 375 5. Venom Glands . . . . . . . . . . 376 6. Quantities and Storage of the Venom 377 7. Toxicity and Human Accidents . . . 377 8. Treatment of Scorpion Envenomation 378
References 378
Table of Contents XIX
B. Chemical and Pharmacologic Aspects of Tityinae Venoms. C.R. DINIZ. With 8 Figures
Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . 379 I. Chemical Properties of Tityinae Venoms . . . . . . . . . 380
1. Purification of Toxic Components from Tityinae Venoms. 380 II. Pharmacology of Tityus Venom Intoxication . . 383
1. Release of Neurotransmitters by Tityus Venom 383 a) Acetylcholine. . . . . . . . . 383 b) Epinephrine and Norepinephrine 386
III. General Effects 391 IV. Conclusion 391
References . . . . . . 391
CHAPTER 15
Chactoid Venoms M. GOYFFON and J. KOVOOR. With 6 Figures
I. Introduction . . . . . . . . . . . . . . . . 395 II. Morphology of the Telson in Chactoids. . . . . . . 396
III. Anatomy and Development of the Venom Glands . . 397 IV. Comparative Histology and Histochemistry of the Venom Glands
in Adults . . . . . . . . . . . . . . . . 398 V. Chemical Composition of Chactoid Venoms. 402
VI. Pharmacology of Chactoid Venoms 406 VII. Experimental Toxicity . . . . . . . . . . 408
VIII. Toxicity for Humans. . . . . . . . . . . 410 IX. Treatment and Prophylaxis of Scorpion Poisoning 412
References . . . . . . . . . . . . . . . . . . . . . 413
CHAPTER 16
Tick Paralysis. M.F. MURNAGHAN and F.J. O'ROURKE. With 3 Figures
A. Introduction . 419 B. Ticks 420
I. The Biology of Ixodid Ticks. 420 II. Classification of Ticks 422
III. Feeding in Ixodid Ticks. 424 IV. Virulence. 425
C. Hosts. 426 I. Humans 426
1. Distribution 426 2. Seasonal Incidence 429 3. Age 429 4. Sex 429
xx Table of Contents
II. Livestock. . . . . . . . 430 1. Natural Occurrence . . 430 2. Experimentally Induced 430
III. Wildlife. . . . . . . 431 IV. Pets . . . . . . . . . . 432
1. Natural Occurrence . . 432 2. Experimentally Induced 433
V. Laboratory Animals 435
D. Symptomology 436 I. Humans . . . . . 436
II. Animals . . . . . 438
E. Mechanism of the Paralysis 439 I. Physiology and Pharmacology of Paralysis due to Dermacentor ander-
soni . . . . . . . . . . . . . 439 1. Neuromuscular Transmission . 439 2. Nerve Conduction. . . . . . 444 3. Central Nervous Transmission 447 4. Miscellaneous Agents . . . . 448
II. Experimental Physiological Findings with Other Ticks Species . 449 1. Ixodes holocyclus . . . . . . . . . . . . . . . . 449 2. Argas (Persicargas) persicus . . . . . . . . . . . 449
III. Identification of the Presence and Removal of the Toxin 449 IV. Immunity. . . . 453 V. Autopsy Findings 454
F. Summary
References .
CHAPTER 17
Toxins of Blattaria. L.M. ROTH and D.W. ALSOP. With 24 Figures
A. Systematics . . . . . . . . . B. Biology .......... . C. Allergy Caused by Cockroaches
1. Contactant Allergens 2. Inhalant Allergens. 3. Injectant Allergens 4. Ingestant Allergens
D. Defensive Glands. . . 1. Type I Glands . 2. Type II Glands. 3. Type III Glands. 4. Type IV Glands. 5. Type V Glands
References
455
458
465 466 467 467 468 469 469
470 471 472 478 481 482
484
Table of Contents
CHAPTER 18
Venoms of Rhyncota (Hemiptera). J. WEATHERSTON and J.E. PERCY
A. Introduction . . . . . . . I. Aquatic Heteroptera . . . . . . . . . .
II. Terrestrial Heteroptera . . . . . . . . .
B. Morphology and Histology of the Scent Glands I. Adult Insects . . . .
II. Larvae ............ .
C. Chemistry of the Secretions . . . . . . I. Structure of Secretion Components.
1. Types of Compounds . . . . . 2. Toxins of Aquatic Heteroptera . 3. Toxins Isolated from Terrestrial Heteroptera 4. Toxins Isolated from Homoptera . . . . .
II. Extraction Methods . . . . . . . . . . . . III. Methods of Purification and Structural Elucidation.
1. Gas Liquid Chromatography 2. Thin-Layer Chromatography 3. Spectroscopic Methods. .
D. Function of Hemipteran Secretions E. Other Glands and Secretions .
References . . . . . . . . . .
CHAPTER 19
Venoms of Coleoptera. J. WEATHERSTON and J.E. PERCY
A. Introduction . . . . . . . . . . . . . . . . B. Morphology and Histology of the Scent Glands
I. Pygidial Gland . 1. Reservoir. . . . . . . 2. Secretory Cells . . . . 3. Fine Structure of Cells. 4. Accessory Gland
II. Thoracic Glands. . . III. Other Glands . . . .
C. Chemistry of the Secretions I. Structure of Secretion Components.
1. Types of Compounds. . 2. Toxins of the Alleculidae 3. Toxins of the Cantharidae 4. Toxins of the Carabidae . 5. Toxins of the Cerambycidae 6. Toxins of the Chrysomelidae
XXI
489 490 490
491 491 494
495 495 496 497 497 497 502 502 504 505 505 505 506
506
511 512 512 512 517 518 519 519 519
520 520 520 521 521 521 526 527
XXII
7. Toxins of the Coccinellidae 8. Toxins of the Dytiscidae. 9. Toxins of the Gyrinidae
10. Toxins of the Meloidae . 11. Toxins of the Silphidae . 12. Toxins of the Staphylinidae 13. Toxins of the Tenebrionidae
II. Extraction Methods . . . . . III. Methods of Purification and Structural Elucidation.
1. Gas Liquid Chromatography ..... 2. Thin-Layer and Paper Chromatography 3. Spectroscopic Methods. . . . . . . .
IV. Biosynthesis. . . . . . . . . . . . . .
D. Biological and Physiological Effects of the Toxins.
References .
CHAPTER 20
Venoms of Lepidoptera. A. DELGADO QUIROZ. With 19 Figures
A. Introduction . . B. Erucism ....
I. Description II. Pathology .
III. Symptomatology . 1. Pathogenesis .
a) Foreign-Body Syndrome. b) Toxic Syndrome .. c) Infectious Syndrome d) Allergic Syndromes
2. Evolution 3. Immunity . . . . .
IV. Diagnosis. . . . . . . 1. Super-Family Papilionoidea (DYAR,1902).
a) Morphidae (KOLLAR, 1850) ..... . b) Nymphalidae (SWAINSON, 1872). . . .
2. Super-Family Bombycoidea (DYAR, 1902) a) Lasiocampidae (HARRIS, 1841) .... b) Thaumetopoeidae (= Notodontidae). . c) Saturniidae (=Hemileucidae) (WALKER, 1855)
3. Super-Family Noctuoidea (MOSHER, 1916) . . . a) Arctiidae (STEPHENS, 1829). . . . . . . . . b) Lymantriidae (= Liparidae) (HAMPSON, 1892) . c) Noctuidae (STEPHENS, 1829) . . . . . . . .
4. Super-Family Zygaenoidea (GRA VENHORST, 1843) a) Cochlidiidae (= Eucleidae) (DYAR, 1898) . b) Megalopigydae (BERG, 1882) ....... .
Table of Contents
527 527 530 533 534 534 535 542 543 543 544 545 545
547
548
555 556 556 558 560 560 560 562 563 563 563 564 564 565 565 565 565 567 568 568 570 570 570 571 571 571 571
Table of Contents
5. The Poison Apparatus. . . . . . . . . . . . . a) Primitive-Type Poison Apparatus (BEYER, 1922) b) Developed-Type Poison Apparatus (FOOT, 1922)
6. Toxicology. . . . . . . a) Obtaining the Poison . . . . . . . . . . . . b) Conserving the Poison. . . . . . . . . . . . c) Pharmacology of the Extracts of Erucic Poisoning
V. Treatment a) Immediate Treatment . . . . . . . . b) Symptomatic and Preventive Treatment c) Treatment of Foreign-Body Syndrome.
VI. Epidemiology . . . . . . . . . . . . 1. Erucism on the American Continent . 2. Erucism on the Continent of Europe. 3. Erucism in Africa, Asia and Australia 4. Epidemiological Structure of Erucism
a) Factors Relating to the Aetiological Agent. b) Factors Relating to Erucic Intoxication . c) Factors Relating to the Environment . .
5. Epidemiological Characteristics of Erucism . 6. Prevention of Erucism . . . . . . . . . .
a) Measures Directed Against the Primary Source.
XXIII
573 575 578 580 580 580 583
584 584 584 585 585 586 588 590 590 590 591 591 592 593 593
b) Measures Directed Against the Means of Transmitting the Poi-sonous Hairs. . . . . . . . . 593
c) Immunization or Desensitization . . . . . . . . . . 593
C. Lepidopterism . .
I. Description . II. Pathology. .
III. Symtomatology 1. Pathogenesis
a) Foreign-Body Syndrome. b) Toxic Syndrome .. c) Infectious Syndrome d) Allergic Syndrome
2. Evolution 3. Immunity . . . . .
IV. Diagnosis. . . . . . . 1. Families with Poisonous Erucae . 2. Zygaenidae Family . . . . . . 3. Satumidae Family (=Hemileucidae) (WALKER, 1855). 4. The Poison Apparatus. . . 5. Toxicology. . . . . . . . .
a) Extraction of the Poison. . b) Conservation of the Poison c) Pharmacology . . . . . .
594
594 594 594 594 595 595 597 597 597 598
598 598 598 598 600 601 601 602 602
XXIV Table of Contents
V. Treatment . . . . . . . . . . 602 a) Immediate. . . . . . . . 602 b) Symptomatic and Preventive 603
VI. Epidemiology and Prevention . . 603 1. South American Lepidopterism 603
a) In the Guianas and Venezuela 603 b) In Peru . . . 603 c) In Brazil. . . . . . . . . . 604 d) In Argentina. . . . . . . . 604
2. Epidemiological Structure of Lepidopterism. 604 a) Factors Relating to the Anthropotoxic Species of the Genus
Hylesia . . . . . . . . . . . . . . . . . . . . . .. 604 b) Factors Relating to the Source of Lepidopteran Intoxication 605 c) Factors Relating to the Environment . . . . . . . 606
3. Prevention of Lepidopterism . . . . . . . . . . . . . .. 606 a) Measures Against the Primary Source of Intoxication . .. 606 b) Measures Against the Means of Transmission of the Poison-
Bearing Flechettes 607
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 607
CHAPTER 21
Venoms of Apidae. R. O'CONNOR and M.L. PECK. With 8 Figures
Introduction. . . . . . . . . . . . . . . . . . . . . . . .
A. Venom Apparatus, Sting Mechanism, and Venom Collection. I. Venom Apparatus
1. Apini .. 2. Bombini ... 3. Meliponini . .
II. Venom Formation . 1. Apini .. 2. Bombini ... 3. Meliponini . .
III. Venom Collection 1. Venom Sac and Gland Extraction. 2. Electrical Excitation of Groups of Bees. 3. Pure Venom from Individual Hymenoptera.
B. Composition of Apis mellifera Venom I. Venom Quantity and Purity.
II. Water Content III. Volatile Components. IV. General Composition. V. Enzymes ..... .
1. Hyaluronidase . . 2. Phospholipase A System 3. Other Enzyme Studies.
613
617 617 617 619 619 619 619 620 620 620 620 621 621
622 622 623 623 624 626 626 628 629
Table of Contents XXV
VI. Large Peptides. . . . . . 630 1. General Characteristics. 630 2. The Melittin Family. . 633 3. Apamin, a Neurotoxin. 634 4. The Mast Cell Degranulating (MCD) Peptide. 635 5. Other Active Peptides . . . . 636
VII. Small Molecules. . . . . . . . . . . 636 1. Some General Considerations. . . . 636 2. Small Peptides and Free Amino Acids 639 3. Biogenic Amines . . . . . . 640 4. Non-Nitrogenous Compounds . . . 640
VIII. Compounds Shown to be Absent . . . 640 IX. Summary of Unresolved Questions on Venom Composition. 641
C. Physiologic Activity of Apis mellifera Venom. 641 I. Direct and Indirect Activity. 641
II. Toxicity Studies . . . . . . . . . . . 645 III. Antigenic Character . . . . . . . . . 646
1. Venom Hypersensitivity and Immunity. 646 2. Antigenic Comparisons among Various Venoms. 647
IV. The Arthritis Question . . 647 1. General Considerations . . . . . 647 2. Possible Mechanisms . . . . . . 647 3. Potentially Interesting Components 648 4. Speculations . . . . . . 648
V. Radioprotective Properties . 648 VI. Other Physiologic Properties 649
D. Venoms of Other Apidae 651 I. Other Apini . 651
II. Bombini 653 Summary. 653
References . . . 653
CHAPTER 22
Venoms of Sphecidae, Pompilidae, Mutillidae, and Bethylidae. W. RATHMAYER. With 8 Figures
A. Introduction . . . . . B. Venoms of Sphecidae .
I. General Aspects . 1. Biological Notes 2. Sting Apparatus and Venom Glands. 3. Stinging of the Prey . . . 4. Venom Actions ..... 5. Effectiveness of Paralysis . 6. Specificity of Venoms . .
661 661 661 661 662 663 665 666 668
XXVI Table of Contents
II. Venoms of Specific Genera . . . . . 672 1. Genus Philanthus . . . . . . . . 672
a) Aspects of Biology and Preying. 672 b) Philanthus triangulum . . . . . 673
IX) Anatomy of the Sting Apparatus. 673 /3) Stinging of the Bee and Action of the Venom 673 y) Mechanism of Action of the Venom 677 <5) Specificity of the Venom 678
2. Genus Sceliphron . . . . . . 680 a) Composition of the Venom 680
C. Venoms of Pompilidae . . . . . . . 680 I. General Aspects. . . . . . . . 680
II. Stinging and Effects of the Venom on Locomotion. 681
D. Venoms of Mutillidae . . . . . . . . . . . . 682 E. Venoms of Bethylidae. . . . . . . . . . . . 682
Appendix 1 : Classification of the Family Sphecidae . 683
References . . . . . . . . . . . . . . . . . . 685
CHAPTER 23
Venoms of Vespidae. H. EDERY, J. ISHAY, S. GITTER and H. JOSHUA. With 30 Figures
A. Introduction . . . . . . . . . . . . . . . . B. Biology of Vespidae. . . . . . . . . . . . .
I. Taxonomy and Geographical Distribution. 1. S teno gastrinae 2. Polistinae 3. Vespinae. .
II. Social Behavior 1. Life Cycle . 2. The Nest Entrance 3. Foraging Behavior. 4. Enemies and Parasites 5. Defensive Behavior . 6. Hornet Traps. . . . 7. Aggressive Behavior-Stinging 8. Aggressiveness of Various Species
C. The Venomous Apparatus ...... . I. The Nonglandular Parts of the Sting Apparatus. Mechanism of
Stinging . . . . . . . . . . . . . . . . II. The Glandular Parts of the Sting Apparatus. . . .
III. Collection of the Venom . . . . . . ..... . IV. Quantity of Venom Collected from a Single Wasp or Hornet
691 691 692 692 693 694 701 701 701 702 704 705 705 705 706
706
706 708 713 715
D. Chemistry of Vespidae Venoms. . . . . . . . . . . . . . . . 715
Table of Contents
1. Physico-Chemical Properties. . . . . II. Components of Vespidae Venoms ..
1. Low-Molecular-Weight Substances a) Acetylcholine. . . . . . . . . b) Histamine . . . . . . . . . . c) Serotonin (5-Hydroxytryptamine, 5-HT) d) Dopamine, Noradrenaline, and Adrenaline. e) Kinins. . . . . . . . . . . . . . . . .
2. High-Molecular-Weight Substances: Enzymes. a) Cholinesterase . . . . . b) Histidine Decarboxylase. . . . . . c) Phospholipases. . . . . . . . . . d) Acid, Alkaline, and Natural DNAses e) Hyaluronidase . . . . . f) Protease. . . . . . . . g) Poly- and Disaccharidases
3. Miscellaneous Substances.
E. Toxicology. . . . . . . . . . . .
l. Systemic Toxicity of V. v. . . . II. Toxicity of V.v. as Related to Ontogenesis
III. Pathological Findings
F. Pharmacology . . . . . . . .
I. Cardiovascular Apparatus.
1. Blood Pressure. . . . 2. Hemolysis. . . . . . 3. Anticoagulant Activity 4. Hyperglycemia. . . . 5. Increase in Permeability of Microcirculation Vessels. 6. Respiratory Apparatus 7. Striated Muscle . . . . . 8. Ocular Irritation . . . . . 9. Isolated Organ Preparations
10. Release of Histamine . . . 11. Pharmacology of Vespidae Kinins
G. Antigenicity and Immunological Aspects.
H. Clinical Aspects of Envenomation. .
1. Incidence of Sting and Dangers II. Symptomatology. . . . .
1. Local Symptoms . . . 2. Generalized Symptoms.
a) Toxic Reactions . b) Allergic Reactions
III. Pathology. IV. Treatment . . . . . .
XXVII
715 715 716 716 718 719 719 720 724 724 724 725 725 725 726 726 726
726
727 728 729
729
731 731 732 739 742 742 743 743 744 744 745 745
748
750
750 752 752 753 753 754 754 755
XXVIII
1. First Aid. . . . . . . . . . . . 2. Emergency Treatment . . . . . . 3. Preventive Therapy-Desensitization
V. Prevention of Stinging VI. Control of Vespidae
J. Concluding Remarks
References . . . . . .
CHAPTER 24
Venoms of Braconidae. R.L. BEARD. With 13 Figures
Introduction.
A. Parasitism by Braconidae I. Stinging and Parasitism.
II. Paralysis of Hosts III. The Preferred Species for Study
B. The Venom Apparatus I. Morphology and Histology
II. Mechanism of Action III. Volume of Ejected Venom
C. Methods of Obtaining Venom for Study. D. Injection and Transport of Venom E. Assays of Venom Activity F. Braconid Venom
I. Chemistry . II. Stability
III. Potency. IV. Production, Storage, and Use of Venom
1. Synthesis. 2. Storage and Volume of Venom Used. 3. Rate of Venom Production.
G. Mode of Action of Venom . I. Effect on the Heart
II. Effect on the Gut III. Effect on Gross Respiration. IV. Effect on Neuromuscular Activity V. Biochemical Studies
H. Host Preferences and Host Sensitivity to Venom 1. Immune Relationships.
References
Table of Contents
755 755 756 756 757
758
759
773
774 774 775 776
777 777 777 778
779 780 781 783 783 783 784 784 785 785 786
786 786 787 788 789 792
794 797
799
Table of Contents XXIX
CHAPTER 25
Venoms and Venom Apparatuses of the Formicidae: Myrmeciinae, Ponerinae, Dorylinae, Pseudomyrmecinae, Myrmicinae and Formicinae. M.S. BLUM and H.R. HERMANN. With 65 Figures
A. Introduction . . . . . . . . . . . . . 801 B. Venoms and Dufour's Gland Constituents 803
I. Myrmeciinae . 803 1. Venom. . . . . . . . . . 803
a) Histamine . . . . . . . 803 b) Smooth Muscle Stimulant 803 c) Hyaluronidase . . . . . 804 d) Direct Hemolytic Factor. 804 e) Phospholipase A . . . . 804 1) Histamine-Releasing Activity . 804
2. Dufour's Gland Constituents 805 II. Ponerinae. . . . . . . . . . 805
1. Venom. . . . . . . . . . 805 2. Dufour's Gland Constituents 805
III. Dorylinae. . . . . 805 1. Venom. . . . . 805
IV. Pseudomyrmecinae . 805 1. Venom. . . . . 805 2. Dufour's Gland Constituents 805
V. Myrmicinae. . . . . . . . . 805 1. Venom. . . . . . . . . . 805 2. Dufour's Gland Constituents 809
VI. Formicinae . . . . . . . . . 810 1. Venom. . . . . . . . . . 810 2. Dufour's Gland Constituents 811
C. Morphology of the Hymenopterous Venom Apparatus 817 D. General Formicid Venom Apparatus . . . . . . . . 817 E. Venom Apparatus Sclerites and their Musculature = General Formicid
Structure . . . . . . . . . . . 819 I. Myrmeciinae and Ponerinae . 827
II. Dorylinae. . . . . 828 III. Pseudomyrmecinae . 830 IV. Myrmicinae. . . . 830 V. Formicinae . . . . 832
F. Abdominal Muscles That Indirectly Affect the Movement of the Venom Apparatus . . . . . . . . . . . 835
G. Muscles of the Venom Apparatus. . . . . . 835 H. Evolution of the Venom Sclerites . . . . . . 835 I. Glands Associated with the Venom Apparatus 837
I. Myrmeciinae, Ponerinae, Dorylinae and Pseudomyrmecinae . 837 II. Myrmicinae . 839
III. Formicinae . . . . . . . . . . . . . . . . . . . . . . 839
xxx Table of Contents
J. Gland Phylogeny ...... . K. Envenomization . . . . . . . L. Mandibular Gland Constituents
I. Ponerinae. . . . . II. Dorylinae. . . . .
III. Pseudomyrmecinae . IV. Myrmicinae .... V. Formicinae . . . .
M. Morphology of the Mandibular Glands N. Metapleural Gland Products . . . . .
I. Myrmicinae. . . . . . . . ..
O. Morphology of the Metapleural Glands P. Natural Products of the Formicidae =An, Overview Q. The Formicid Venom Apparatus = Phyletic Implications
I. Relationship to Wasps II. Formicinae . . . .
III. Pseudomyrmecinae. . IV. Myrmicinae ..... V. Ponerinae, Dorylinae, Cerapachyinae .
VI. Old and New World Dorylinae
References . . . . . . . . . . . . . . . .
CHAPTER 26
840 843 844 844 845 846 846 849
851 853 853
854 854 857 857 859 859 860 860 860
860
Venom and Venom Apparatuses of the Formicidae: Dolichoderinae and Aneuretinae. M.S. BLUM and H.R. HERMANN, JR. With 11 Figures
A. Introduction and Taxonomy . . . . . . . . . . . . . . . . . . . . 871 B. Dolichoderine Natural Products and Anatomy of the Venom Apparatus. 872
I. Mandibular Gland Constituents . . . . . . . . . . . . . . . . 872 II. Poison Gland Constituents . . . . . . . . . . . . . . . . . . 873
III. Anatomy of the Venom Apparatus Components in the Dolichoderi-nae . . . . . . . . . . . 874·
IV. Anatomy of Venom Sclerites 875 V. Dufour's Gland Constituents 877
VI. Anatomy of Dufour's Gland 880
C. Anal Gland Constituents I. Anatomy of the Anal Glands in the Dolichoderinae
II. Biological Activities of Dolichoderine Anal Gland Products.
D. Dolichoderine Natural Products=An Overview. . ...... . E. Phyletic Implications Based on Venom Slerites and Associated Glands.
References .
Author Index
Subject Index
880 884 885
886 890
891
895
. 940
List of Contributors
D.W. ALSOP, Dr., Department of Biology, Queen's College, City University of New York, Flushing, NY 11367, USA
R.L. BEARD, Dr., 864 Mountain Road, Cheshire, CT 06410, USA
S. BETTINI, Professor Dr., Istituto Superiore di Sanita, Viale Regina Elena 299, 1-00161 Rome, Italy
M.S. BLUM, Professor, Department of Entomology, University of Georgia, Athens, GA 30602, USA
P.M. BRIGNOLI, Professor Dr., Universita degli Studi di L' Aquila, Istituto di Zoologia, Piazza Regina Margherita 7, 1-67100 L'Aquila, Italy
W. BUCHERL, Professor Dr., Rua Sagarana 152, Vila Madalena, 0544 Sao Paulo, Brazil
A. DELGADO QUIROZ, Dr., Instituto de Medicina Tropical, "Daniel A. Carrion ", Universidad Nacional Mayor de San Marcos, Casilla 10138, Lima I, Peru
C.R. DINIZ, Dr., Ministerio da Educac;ao E Cultura, Universidade Federal De Minas Gerais, CPo 2486, Instituto De Ciencias Bio16gicas, Dept. Bioquimica, 30.000 Belo Horizonte, M.G., Brazil
H. EDERY, Professor, Department of Physiology and Pharmacology, Sackler School of Medicine, Tel-Aviv University, Ramat Aviv, Israel
P. EFRATI, Professor, Kaplan Hospital, Rehovot, Israel
T. EISNER, Professor, Section of Neurobiology and Behavior, Cornell University, 139 Langmuir Laboratory, Ithaca, NY 14853, USA
S. GITTER, Professor, Department of Physiology and Pharmacology, Sackler Scholl of Medicine, Tel-Aviv University, Ramat-Aviv, Israel
M. GOYFFON, Dr., Museum National d'Histoire Naturelle, Laboratoire d'Etudes et de Recherches sur les Animaux Irradies C.R.S.S.A. - C.N.R.S., 57 Rue Cuvier, F-75 Paris 5e , France
M.R. GRAY, Dr., The Australian Museum, P.O. Box A 285, Sydney, N.S.W. 2000, Australia
Y. HASHIMOTO, Professort, The University of Tokyo, Faculty of Agriculture, Laboratory of Marine Biochemistry, Bunkyo-ku, Tokyo, Japan
XXXII List of Contributors
H.R. HERMANN, Professor, Department of Entomology, University of Georgia, Athens, GA 30602, USA
KAREN HICKS, Dr., Section of Neurobiology and Behavior, 139 Langmuir Laboratory, Cornell University, Ithaca, NY 14853, USA
J. ISHAY, Dr., Department of Physiology and Pharmacology, Sackler School of Medicine, Tel-Aviv University, Ramat-Aviv, Israel
H. JOSHUA, Dr., Clinical Laboratory, Beilinson Medical Center, Petah Tikva, Israel
S. KONOSU, Dr., The University of Tokyo, Faculty of Agriculture, Laboratory of Marine Biochemistry, Bunkyo-ku, Tokyo 113, Japan
JACQUELINE KOVOOR, Universite Pierre-et-Marie Curie, Laboratoire d'Evolution, 105, Bd. Raspail, F-75006 Paris, France
G. LEVY, Dr., Department of Entomology and Venomous Animals, The Hebrew University, Jerusalem, Israel
Z. MARETIC, Dr., Medicinski centar, Pula, Jugoslavia
M. MAROLI, Dr., Laboratorio di Parassitologia, Istituto Superiore di Sanita, Viale Regina Elena 299, 1-00161 Rome, Italy
J. MEINWALD, Professor Dr., Department of Chemistry, Cornell University, Spencer T. Olin Laboratory, Ithaca, NY 14853, USA
A. MINELLI, Professor Dr., Istituto di Biologia Animale, Universita degli Studi, Via Poredan 10, 1-35100 Padova, Italy
F. MIRANDA, Dr., Laboratoire de Biochimie, Faculte de Medecine-Secteur Nord, F-13326 Marseille Cedex 3, France
M.F. MURNAGHAN, Professor, Department of Physiology, Earlsfort Terrace, University College, Dublin, Ireland
R. O'CONNOR, Dr., Chemistry Department, Texas University, College Station, TX 77843, USA
F.J. O'ROURKE, Dr., Department of Zoology, University College, Lee Maltings Prospect Row, Cork, Ireland
M.L. PECK, Ass. Professor, Department of Chemistry, Texas A+M University, College of Science, College Station, TX 77843, USA
J.E. PERCY, Dr., Canadian Forestry Service, Insect Pathology Research Institute, P.O. Box 490, Sault Ste. Marie, Ontario P6A 5M7, Canada
F.A. PEREIRA-LIMA, Dr., Servico de Fisiologia do Instituto Butantan, Caixa Postal 65, Sao Paulo, Brazil
W. RATHMAYER, Professor, Fachbereich Biologie, Universitiit Konstanz, Postfach 733, 7750 Konstanz, Germany
List of Contributors XXXIII
H. ROCHAT, Dr., Laboratoire de Biochimie, Faculte de Medecine - Secteur Nord, F-13326 Marseille, Cedex 3, France
L.M. ROTH, Dr., 81 Brush Hill Road, Sherborn, Massachusettes, USA 01770
S. SCHENBERG, Professor, Servico de Fisiologia, Instituto Butantan, Caixa Postal 65, Sao Paulo, Brazil
H. SCHENONE, Dr., Instituto de Parassitologia, Universidad de Chile, Cassila 9183, Santiago de Chile, Chile
A. SHULOV, Professor Dr., Department of Entomology and Venomous Animals, The Hebrew University, Jerusalem, Israel
H.L. STAHNKE, Professor, Poisonus Animals Research Laboratory, Department of Zoology, Arizona State University, Tempe, Arizona 85281, USA
G. SUAREZ, Dr., Department of Medicine, Albert Einstein College of Medicine, 1300 Morris Park Avenue, Bronx, NY 10461, USA
S.K. SUTHERLAND, Dr., Department of Immunological Research, Commonwealth Serum Laboratories, 45 Poplar Rd., Parkville, Melbourne 3052, Australia
J. WEATHERSTON, Dr., Forest Pest Management Institute, 1219 Queen Street East, Sault Ste. Marie, Ontario, Canada
E. ZLOTKIN, Dr., The Hebrew University of Jerusalem, Department of Entomology and Venomous Animals, Jerusalem, Israel
