Snake Venoms and Envenomations
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SNAKE VENOMS AND ENVENOMATIONS Jean-Philippe Chippaux Translation by F. W. Huchzermeyer KRIEGER PUBLISHING COMPANY Malabar, Florida 2006 Original French Edition Venins des"pent et enuenimations 2002 Translated from the original French by F. W. Huchzermeyer Copyright © English Edition 2006 by IRD Editions. France Original English Edition 2006 Printed and Published by KRIEGER PUBLISHING COMPANY KRIEGER DRIVE MAlABAR. FLORIDA 32950 Allrights reserved. No part of this book may be reproduced in any form or by any means. electronic or mechanical. including information storate and retrieval systems without permission in writing from the publisher. No liabilityisassumed with respect to the useofthe inftrmation contained herein. Printed in the United States of America. Library of Congress Cataloging-in-Publication Data FROM A DECLARATION OF PRINCIPLES JOINTLY ADOPTED BY A COMMITTEE OF THE AMERICAN BAR ASSOCIATION AND A COMMITTEE OF PUBLISHERS: This publication is designed to provide accurqate and authoritative information in regard to the subject matter covered. It is sold with the understanding that the publisher is not engaged in rendering legal, accounting. or other professional service. If legal advice or other expert assistance is required. the services of a competent professional person should be sought. Chippaux, Jean-Philippe. [Venins de serpent et envenirnations, English] Snake venoms and envenomations I Jean-Philippe Chippaux ; translation by F.W. Huchzermeyer. p. cm. Includes bibliographical references and index. ISBN 1-57524-272-9 (hbk. : a1k. paper) 1. Venom. 2. Snakes. 3. Toxins. I. TIde. QP235.C4555 2006 615.9'42---<ic22 2005044486 10 9 8 7 6 5 4 3 2 Contents Glossary v Preface vii Introduction ix PART I ZOOLOGY 1. The Zoology ofSnakes 3 Palaeontology 3 Anatomy 18 The Systematicsofthe Snakes 24 PART 11 VENOMS 2. The Venom Apparatus 47 The General Evolution ofthe Venom Apparatus 47 The Anatomy ofthe Venom Apparatus 51 The Composition ofthe Venom 54 3. The Toxicology ofthe Venoms 75 Measuring Toxicity 75 Effectson Cells 78 Effects on the Nervous System 81 Effectson the Cardio-vascular System 89 Effectson Haemostasis 91 Necrotizing Activity 107 Toxicokinetics 112 The Utilization ofVenoms in Research and Therapeutics 115 111 IV Contents 4. Antidotes and Immunotherapy 125 Traditional Medicine 125 Principles and Mechanisms ofImmunotherapy 136 Medical Treatment 161 Vaccination 164 PART III ENVENOMATIONS 5. The Epidemiology ofEnvenomations 169 The Ecology ofSnakes 169 Human Behavior 182 The Epidemiology ofBites 185 6. Clinic and Treatment ofEnvenomations 211 Symptomatology and Clinical Pathology ofEnvenomations 211 TheTreatment ofEnvenomations 228 Evaluation ofthe Availability ofAntivenins and the Improvement oftheir Distribution 239 Prevention ofEnvenomations 242 Appendix 1 Practical Measuring ofthe LD lQ by the Spearrnan- Karber Method 247 Appendix 2 Symptomatic Action ofCertain Plants 249 Appendix 3 Antivenom Producers and Their Products 257 Appendix 4 French Legislation Regarding the Keeping ofVenomous Animals 271 Bibliography 275 Index 281 Glossary Absolute density: The real number ofindividuals per unit ofsurface or volume after an exhaustive count ofall individuals Abundance: The number ofindividuals belonging to one population Action potential: Electrical wave moving along a nerve fibre and provoking a muscle contraction AgIyphous: Maxilla with full teeth not linked to salivary or venom glands- fang-less Alethinopbidian: Group ofmodern shakes, ethymologically "true snakes" Apodal: without limbs Apparent density: Relative number ofindividuals per unit ofsurface or volume as function ofa collection method or collector's activity Autochthonous: Oflocal origin Dentition: The number and distribution ofteeth on the jaws; this is species specific Diasteme: Free space on the maxilla separating two series ofteeth Duvemoy: Sero-mucous salivary gland which in the colubrids produces a toxic secretion, a precursor ofthe venom gland Effector site: Precise site on a molecule where an activity is situated Endemic: Naturally occurring in an area or region Endothelium: Layer ofcells covering the inside ofblood vessels preventing leakage and setting into motion the coagulation in the case ofinjury Epitope: The exact site ofa molecule to which antibodies bind Evolutionary radiation: The appearance under environmental pressure of new zoological forms from a common ancestor Fibrillation: Repeated short muscular contraction oflow amplitude Haemostasis: Phenomenon or action aimed at stopping a hemorrhage Incidence: Frequency ofan event per period oftime (usually one year); in the case ofenvenomations the incidence refers specifically to all bites irrespective of their severity Jacobson's organ: Sensory organ ofsnakes associated with the sense ofsmell vomero-nasal organ Labial: Lip region; the labials are he scales adjacent to upper and lower lips v VI Glossary Lethality: The frequency ofdeaths caused by a given condition Lorea1: The lateral portion ofthe head between to eye and the nostril; the loreal shield is the shield covering this region without contact with eye or nostril Monophyletic: Zoological groups sharing the same ancestor Morbidity: The frequency ofa disease per period oftime (generally one year); in the case ofenvenomations the morbidity represents the symptomatic bites and thus a fraction of the incidence Naso-vomeral organ - SeeJacobson's organ Opisthodontous: Presence ofa tooth at the back of the maxilla and separated from the preceding ones by a diasteme Opisthoglyphous: Presence ofa poison fang at the back ofthe maxilla and separated from the preceding ones by a diasterne, back-fanged Pleurodontous: Sentition containing different types ofteeth Population: All the individuals belonging to the same species and living in a given geographical zone Prevalence: Number ofcases diagnosed at a given moment expressing the present frequency ofan event Proterodontous: Presence ofa larger tooth at the front end ofthe maxilla Proteroglyphous: Tubular tooth or tooth with an open canal carried on a generally short fIXed or slightly mobile maxilla, front-fanged Purpura: Eruption of red blotches on the skin due to blood seeping out of the capillaries due to the rupture ofthe blood vessels or to a hemorrhage problem Rhodopsin: Retina pigment needed for colour vision Scolecophidian: Group of primitive still living snakes; ethymologically "worm snakes" Solenoglyphous: Tubular fang or one with an open canal situated on a short and mobile maxilla Symphysis: Fixed or slightly mobile articulation consisting ofcartilage and elastic tissue Territory:The space in which an animal moves and where it completes its principal activities in the rhythm ofits existence (feeding, mating, egg laying or giving birth), lebensraum Vitellogenesis: Deposition offat reserves by the females in the ovarian follicles before ovulation Preface While the mythical representation of snakes in all cultures originated from an ignorance of their true nature, it was also responsible for the delay with which sci entists took up the study of these very peculiar animals: Often the legends sub stituted observation and experimentation for their description and explication. Nonetheless, with the exception ofa few countries without venomous species, poisonous snakebites are feared throughout the whole world and pose a real pub lic health problem in the tropical regions, in particular in developing countries. There are approximately 2700 described species ofsnakes ofwhich 500 are ven omous and potentially dangerous to people in varying degrees. The majority of bites result from the unexpected but not purely accidental encounter between human and snake and in most cases the latter only defends itself against what it considers to be an aggressor. Epidemiological studies aim at defining the causes and origins of these encounters. They need to be based on zoological research, which helps to identify the species involved, to study their ecology and this could lead to suggestions how to prevent these bites where possible. In addition the specificity of the venoms has been the subject ofdetailed biochemical and toxi cological studies which have allowed us to understand the diversity ofsymptoms observed in cases ofsnakebite and to suggest specific treatments. Every year there are 6 million snakebites in the entire world, half of them with. subsequent clinical problems ofvarying severity.The annual number offatal cases is estimated to be 125,000, most ofthem in Africa and Asia, namely in poor coun tries where generally treatment is inaccessible. Most snakebites happen in rural zones during agricultural activities. In contrast, in developed countries where the dangerous activities are more rare because of the mechanization of agriculture, accidental bites have appeared over the last few years which occur during recre ational activities, as well as "illegitimate" bites due to handling ofpet snakes, some ofwhich are exotic and particularly dangerous. In all cases the envenomations are really difficult to take care ofand to treat. vu Vll1 Preface However, even beyond what one could consider a simple defense strategy, the study ofthe biology ofsnakes, of the composition and mode ofaction of the ven oms has opened new and fascinating research perspectives regarding the molecu lar structure of numerous membrane receptors, the intimate mechanisms of intercellular relations or the discovery ofnew effective drugs against hitherto untreatable diseases. This book offers a synthesis ofrecent