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Parasitology JWST138-Fm JWST138-Gunn February 21, 2012 16:59 Printer Name: Yet to Come P1: OTA/XYZ P2: ABC JWST138-fm JWST138-Gunn February 21, 2012 16:59 Printer Name: Yet to Come P1: OTA/XYZ P2: ABC Parasitology JWST138-fm JWST138-Gunn February 21, 2012 16:59 Printer Name: Yet to Come P1: OTA/XYZ P2: ABC Parasitology An Integrated Approach Alan Gunn Liverpool John Moores University, Liverpool, UK Sarah J. Pitt University of Brighton, UK Brighton and Sussex University Hospitals NHS Trust, Brighton, UK A John Wiley & Sons, Ltd., Publication JWST138-fm JWST138-Gunn February 21, 2012 16:59 Printer Name: Yet to Come P1: OTA/XYZ P2: ABC This edition first published 2012 © 2012 by by John Wiley & Sons, Ltd Wiley-Blackwell is an imprint of John Wiley & Sons, formed by the merger of Wiley’s global Scientific, Technical and Medical business with Blackwell Publishing. Registered Office John Wiley & Sons Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial Offices 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 111 River Street, Hoboken, NJ 07030-5774, USA For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell. The right of the author to be identified as the author of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data Gunn, Alan. Parasitology : an integrated approach / Alan Gunn and Sarah J. Pitt. p. ; cm. Includes bibliographical references and index. ISBN 978-0-470-68424-5 (cloth) – ISBN 978-0-470-68423-8 (pbk.) I. Pitt, Sarah J. II. Title. [DNLM: 1. Host-Parasite Interactions. 2. Parasites–physiology. 3. Parasitic Diseases. QY 45] 616.96–dc23 2011043529 A catalogue record for this book is available from the British Library. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. Set in 10.5/12.5pt Times by Aptara Inc., New Delhi, India First Impression 2012 JWST138-fm JWST138-Gunn February 21, 2012 16:59 Printer Name: Yet to Come P1: OTA/XYZ P2: ABC None of us truly live alone JWST138-fm JWST138-Gunn February 10, 2012 9:40 Printer Name: Yet to Come P1: OTA/XYZ P2: ABC Contents Preface xiii 1 Animal associations 1 1.1 Introduction 1 1.2 Animal associations 1 1.2.1 Symbiosis 2 1.2.2 Commensalism 5 1.2.3 Phoresis 5 1.2.4 Mutualism 6 1.2.5 Parasitism 7 1.2.6 Intra-specific parasites 8 1.2.7 Parasitoids 9 1.2.8 The concept of harm 10 1.3 Parasite hosts 11 1.3.1 Protozoa and helminths as hosts 11 1.3.2 Classes of hosts for parasites 12 1.4 The co-evolution of parasites and their hosts 13 1.4.1 Evolutionary relationships between host and parasite 14 1.4.2 Parasites and the evolution of sexual reproduction 15 1.5 Parasitism as a ‘lifestyle’: advantages and limitations 16 1.5.1 Main advantages of a parasitic lifestyle 17 1.5.2 Main limitations of a parasitic life style 17 1.6 The economic cost of parasitic diseases 18 1.6.1 Economic consequences of parasitic diseases of humans 18 1.6.2 Economic consequences of parasitic diseases of domestic animals 19 1.6.3 Estimating the costs of morbidity due to disease 19 1.6.4 Economic consequences of parasitic diseases of wildlife 20 1.7 Why parasitic diseases remain a problem 21 1.8 Taxonomy 24 1.8.1 The binomen system 25 Questions 27 2 Parasitic protozoa, fungi and plants 28 2.1 Introduction 28 2.2 Parasitic protozoa 28 2.2.1 Kingdom Protista 28 JWST138-fm JWST138-Gunn February 10, 2012 9:40 Printer Name: Yet to Come P1: OTA/XYZ P2: ABC viii CONTENTS 2.3 Phylum Rhizopoda 29 2.3.1 Genus Entamoeba 29 2.3.2 Other species of pathogenic amoebae 33 2.4 Phylum Metamonada 34 2.4.1 Order Diplomonadida 34 2.4.2 Order Trichomonadida 37 2.5 Phylum Apicomplexa 40 2.5.1 Genus Plasmodium 42 2.5.2 Plasmodium life cycle 43 2.5.3 Genus Theileria 45 2.5.4 Genus Babesia 46 2.6 Subclass Coccidiasina 50 2.6.1 Suborder Eimeriorina 50 2.6.2 Isospora group 51 2.6.3 Genus Cyclospora 52 2.6.4 Family Sarcocystidae 53 2.6.5 Genus Toxoplasma 54 2.6.6 Genus Neospora 58 2.6.7 Family Cryptosporidiidae 60 2.7 Phylum Kinetoplastida 62 2.7.1 Genus Leishmania 63 2.7.2 Leishmania life cycle 66 2.7.3 Genus Trypanosoma 70 2.8 Phylum Chlorophyta 81 2.8.1 Genus Prototheca 82 2.9 Kingdom fungi 83 2.9.1 Microsporidia 83 2.10 Kingdom plantae 85 Questions 85 3 Helminth parasites 86 3.1 Introduction: invertebrate taxonomy 86 3.2 Phylum Platyhelminthes 87 3.3 Class Trematoda 87 3.3.1 Family Fasciolidae 89 3.3.2 Family Cathaemasiidae: Genus Ribeiroia 93 3.3.3 Family Dicrocoeliidae 95 3.3.4 Family Opisthorchiformes 96 3.3.5 Family Paragonomidae 98 3.3.6 Family Schistosomatidae 99 3.4 Class Cestoda 103 3.4.1 Order Pseudophyllidea/Diphyllobothriidea 103 3.4.2 Order Cyclophyllidea 104 3.4.3 Family Taeniidae 105 3.4.4 Family Anoplocephalidae 110 3.5 Phylum Acanthocephala 112 3.6 Phylum Nematoda (Nemata) 114 3.6.1 Class Enoplea 117 3.6.2 Class Rhabdita 121 3.6.3 Family Onchocercidae 129 3.6.4 Family Dracunculidae 132 Questions 135 JWST138-fm JWST138-Gunn February 10, 2012 9:40 Printer Name: Yet to Come P1: OTA/XYZ P2: ABC CONTENTS ix 4 Arthropod parasites 137 4.1 Introduction 137 4.2 Phylum Chelicerata 138 4.2.1 Family Demodicidae 139 4.2.2 Family Sarcoptidae 140 4.2.3 Family Psoroptidae 143 4.2.4 Suborder Ixodida 144 4.2.5 Family Argasidae 145 4.2.6 Family Ixodidae 146 4.2.7 Tick paralysis 147 4.3 Phylum Crustacea 148 4.3.1 Subclass Copepoda 148 4.3.2 Infra-Class Cirripedia 150 4.3.3 Subclass Branchiura 150 4.3.4 Subclass Pentastomida – tongue worms 151 4.4 Sub-phylum Hexapoda 153 4.4.1 Order Phthiraptera (lice) 155 4.4.2 Order Siphonaptera (fleas) 159 4.4.3 Order Diptera (true flies) 162 4.4.4 Suborder Nematocera 162 4.4.5 Suborder Brachycera 163 4.4.6 Family Calliphoridae 166 4.4.7 Genus Chrysomya 168 4.4.8 Genus Cochliomyia 168 4.4.9 Genus Auchmeromyia 169 4.4.10 Genus Cordylobia 170 4.4.11 Family Sarcophagidae 170 4.4.12 Family Oestridae 171 4.4.13 Subfamily Gasterophilinae 173 4.4.14 Subfamily Hypodermatinae 174 4.4.15 Subfamily Cuterebrinae 176 4.4.16 Family Streblidae 177 4.4.17 Family Nycteribiidae 178 Questions 178 5 Parasite transmission 180 5.1 Introduction 180 5.2 Contaminative transmission 181 5.3 Transmission associated with reproduction 184 5.3.1 Sexual transmission 184 5.3.2 Transmission within the gametes 187 5.3.3 Congenital transmission 188 5.4 Autoinfection 189 5.5 Nosocomial transmission 190 5.6 Active parasite transmission 191 5.7 Hosts and vectors 192 5.7.1 Paratenic hosts 192 5.7.2 Intermediate hosts 193 5.7.3 Vectors 194 5.8 Host factors 196 5.8.1 Host identification 196 5.8.2 The influence of host behaviour on parasite transmission 197 JWST138-fm JWST138-Gunn February 10, 2012 9:40 Printer Name: Yet to Come P1: OTA/XYZ P2: ABC x CONTENTS 5.9 Co-transmission and interactions between infectious agents 199 5.10 How religion can influence parasite transmission 202 5.11 The influence of war on parasite transmission 204 5.12 The influence of parasites on host behaviour 205 5.13 Environmental factors 207 5.13.1 Natural environmental variables 207 5.13.2 Pollution 207 5.13.3 Global warming 209 Questions 211 6 Immune reactions to parasitic infections 212 6.1 Introduction 212 6.2 Invertebrate immunity 213 6.3 Vertebrate immunity 215 6.3.1 Innate immunity 215 6.3.2 Adaptive immunity 218 6.3.3 Cell-mediated immunity 220 6.4 Innate immunity to parasitic infection 221 6.4.1 Physical factors 221 6.4.2 Chemical and microbial factors 222 6.4.3 The acute inflammatory response 223 6.4.4 Cell-mediated immunity 225 6.5 Adaptive immunity 226 6.5.1 Avoiding the host immune response 227 6.5.2 Depression of the immune system 232 6.6 Immunity to malaria 233 6.7 Schistosoma mansoni and Hepatitis C virus interactions 237 6.8 HIV-AIDS and parasitic disease 238 6.8.1 Parasites and the transmission of HIV 239 6.8.2 Parasite-HIV co-infections 240 6.8.3 Leishmania-HIV co-infections 240 6.8.4 Malaria–HIV co-infections 242 6.8.5 Toxoplasma–HIV co-infections 243 6.8.6 Microsporidia–HIV co-infections 243 Questions 243 7 Pathology 245 7.1 Introduction 245 7.2 Factors that influence pathogenesis 245 7.2.1 Host factors that influence pathogenesis 245 7.2.2 Parasite factors that influence pathogenesis 246 7.3 Mechanisms by which parasites induce pathology 247 7.3.1 Direct damage 248 7.3.2 Indirect damage 249 7.4 Types of pathology 250 7.4.1 Abortion and obstetric pathology 250 7.4.2 Anaemia 251 7.4.3 Anorexia 253 7.4.4 Apoptosis 253 7.4.5 Calcification 254 JWST138-fm JWST138-Gunn February
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