Human Parasitology
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HUMAN PARASITOLOGY FOURTH EDITION BURTON J. BOGITSH,PHD CLINT E. CARTER,PHD THOMAS N. OELTMANN,PHD AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Academic Press is an imprint of Elsevier 225 Wyman Street, Waltham, MA 02451, USA The Boulevard, Langford Lane, Kidlington, Oxford, OX5 1GB, UK Ó 2013 Elsevier Inc. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the Publisher. Details on how to seek permission, further information about the Publisher’s permissions policies and our arrangements with organizations such as the Copyright Clearance Center and the Copyright Licensing Agency, can be found at our website: www.elsevier.com/permissions This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be noted herein). Notices Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding, changes in research methods, professional practices, or medical treatment may become necessary. Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any information, methods, compounds, or experiments described herein. In using such information or methods they should be mindful of their own safety and the safety of others, including parties for whom they have a professional responsibility. To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions, or ideas contained in the material herein. Library of Congress Cataloging-in-Publication Data Bogitsh, Burton J. (Burton Jerome) Human parasitology / Burton J. Bogitsh, Clint E. Carter. e 4th ed. p. cm. Includes bibliographical references and index. ISBN 978-0-12-415915-0 (alk. paper) 1. Medical parasitology. 2. Parasitic diseases. I. Carter, Clint E. (Clint Earl) II. Title. QR251.B47 2013 616.9’6–dc23 2011053044 British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library. For information on all Academic Press publications visit our website at http://store.elsevier.com Printed in the United States of America 12 13 14 15 10 9 8 7 6 5 4 3 2 1 Dedicated to my past and present inspirations, Mafoi and Teresa -BJB and to Patty (CEC) and Peggy (TNO) Preface Several years ago we recognized the need subsequently included in the sections in for a textbook of human parasitology which host immune responses are discussed designed specifically for premedical, medical relative to the parasite(s) in question. technology, and biology students who We still feel that it is important that the required basic knowledge of the biology of biological aspects of parasites be considered. parasitism, and we began to consider collab- To this end, we have preserved the opening orating on such a text. As the idea germi- chapter to each major group of parasites nated, we decided that, while emphasizing and kept the designation as “General Char- the medical aspects of the topic, the book acteristics.” In these introductory chapters should incorporate sufficient functional we deal with those characteristics that are morphology, physiology, biochemistry, and universal within the group as a whole. We immunology to enhance appreciation of the deal with the more specialized aspects of diverse implications of parasitism. It would individual parasites in their proper context. also explore the potential of certain parasites Ample light and electron micrographs serve for producing morbidity and mortality and to illustrate the various points of the text. would present available data regarding the An abridged classification of parasites modus operandi of certain modern chemother- covering the major nomenclature has been apeutic agents. Through considerable discus- placed at the end of each chapter dealing sion and several revisions of the manuscript, with the general characteristics of each the first edition of Human Parasitology group. At the end of most chapters is evolved. One of the goals for Human Parasi- a section entitled “Selected Readings.” We tology was that it would serve as a bridge have attempted to list a few significant publi- between classical clinical parasitology texts cations that present some in-depth informa- and the more traditional encyclopedic, tion on selected topics pertinent to the advanced treatises that include in-depth topics discussed in the chapter. This section consideration of biochemistry and immu- is not intended to present an updated litera- nology as well as the inclusion of those para- ture for research or graduate studies. sites that infect nonhuman hosts (e.g., However, the reader will find some classical strigeids of fishes, amphibians, and birds). reports as well as more recent citations. As described earlier, the subject of oppor- The field of chemotherapy is advancing at tunistic parasites was expanded in the third such a rapid pace that new drugs and regi- edition and with this expansion the section mens become available almost continually. on immunology was accordingly expanded Consequently, the sections dealing with and updated. In this edition, we have information about current drugs of choice expanded and simplified the topics of innate as well as regimens will possibly become and acquired immunity emphasizing the obsolete more rapidly than other sections role that toll-like receptors play in the innate of the book. Nevertheless, in the individual immune response. This information was sections and at the end of the book, not xv xvi PREFACE only have we included specific, currently new material gleaned from genomic studies prescribed chemotherapeutic drugs and such as identifying differences in virulence regimens but we have updated, as of the of different strains of certain parasites such time of this writing, the newest changes in as Toxoplasma or identifying targets for new these protocols. For instance, we have drugs in Cryptosporidium. A number of other included facts about Artemisinin as an anti- changes have been incorporated in the text malarial. We also discuss how the use of such as standardization of illustrations and combinations of drugs proved useful in section headings that improve the aesthetics combating onchocerciasis. of the text. In conclusion, we believe this Because the current edition, as well as edition will appeal to those students inter- previous editions, is designed for a one- ested not only in the medical aspects of para- semester course, the material is somewhat sitology but also to those who require a solid condensed, particularly in the areas of phys- foundation in the biology of parasites in iology and biochemistry. To stay current with order to further their studies in a graduate research in these fields, we have included or a professional school of their choice. CHAPTER 1 Symbiosis and Parasitism OUTLINE Definitions 2 Diagnosis 9 Commensalism 2 Factors Influencing Prevalence 9 Phoresis 2 Unsanitary Living Conditions 10 Parasitism 2 Disease Control and Treatment 10 Mutualism 4 Poor Nutrition 10 Health Education 10 Ecological Aspects of Parasitism 5 Regional and Ethnic Customs 10 Medical Implications 7 Climatic Conditions 11 Control Impediments 8 Opportunistic Parasitism 11 Intermediate Hosts 8 Evolution of Parasitism 11 Vectors 8 Resistance and Resurgence 9 Parasitology, the study of parasites and their relationships to their hosts, is one of the most fascinating areas of biology. The study of parasitism is interdisciplinary, encompassing aspects of systematics and phylogeny, ecology, morphology, embryology, physiology, biochemistry, immunology, pharmacology, and nutrition, among others. Newly developed techniques in biochemistry and cellular and molecular biology have also opened significant new avenues for research on parasites. Not only does parasitology touch upon many disciplines, but the varied nature of para- sites renders their study multifaceted. While it is entirely proper to classify many bacteria and fungi and all viruses as parasites, parasitology has traditionally been limited to parasitic protozoa, helminths, arthropods, and those species of arthropods that serve as vectors for parasites. It follows, then, that parasitology encompasses elements of protozoology, helmin- thology, entomology, and acarology. The World Health Organization has proclaimed that, of the six major unconquered human tropical diseases, fivedschistosomiasis, malaria, filariasis, Human Parasitology, Fourth Edition 1 DOI: 10.1016/B978-0-12-415915-0.00001-7 Ó 2012 Elsevier Inc. All rights reserved. 2 1. SYMBIOSIS AND PARASITISM African trypanosomiasis, and leishmaniasisdare parasitic in the traditional sense. Leprosy, the sixth major disease, is caused by a bacterium. DEFINITIONS The complexity of the host-parasite relationship has often led to misunderstandings of the precise nature of parasitism. In order to avoid such misperceptions, researchers have devised the following concepts to distinguish among the several types of associations involving het- erospecific organisms. Any organism that spends