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Veterinary Microbiology VETERINARY MICROBIOLOGY Dwight C. Hirsh, DVM, PhD Professor, Department of Pathology, Microbiology, and Immunology Chief of Service, Clinical Microbiology Laboratory ‘Veterinary Medical Teaching Hospital School of Veterinary Medicine University of California lifornia Davis, C Yuan Chung Zee, DVM, PhD Professor, Department of Pathology, Microbiology, and Immunology School of Veterinary Medicine University of California Davis, California b Blackwet Selene © 1999 by Blackwell Science, Inc. Editorial Offices: Commerce Place, 350 Main Street, Malden, Massachusetts 02148, USA Osney Mead, Oxford OX2 OEL, England 25 John Street, London WCIN 2BL, England 23 Ainslie Place, Edinburgh EH3 6AJ, Scotland 54 University Street, Carlton, Victoria 3053, Australia Other Editorial Offices: Blackwell Wissenschafts-Verlag GmbH, Kurfiirstendamm, 57, 10707 Berlin, Germany Blackwell Science KK, MG Kodenmacho Building, 7-10 Kodenmacho Nihombashi, Chuo-ku, Tokyo 104, Japan Distributors: USA Blackwell Science, Inc. ‘Commerce Place 350 Main Street Malden, Massachusetts 02148 (Telephone orders: 800-215-1000 or 781-388-8250; fax orders: 781-388-8270) Canada Login Brothers Book Company 324 Saulteaux Crescent Winnipeg, Manitoba, R3 372 (Telephone orders: 204-224-4068) Australia Blackwell Science Pty, Ltd. 54 University Street Carlton, Victoria 3083 (Telephone orders: 03-9347-0300; fax orders: 03-9349-3016) Outside North America and Australia Blackwell Science, Ltd. ‘c/o Marston Book Services, Ltd. P.O. Box 269 Abingdon Oxon OX14 4YN England (Telephone orders: 44-01235-465500; fax orders: 44-01235-465555) Al rights reserved. No part of this book may be reproduced in any form or by any electronic or mechanical means, including information storage and retrieval systems, without permission in writing from the publisher, except by a reviewer who may quote brief passages in a review. ‘Acquisitions: Nancy Hill-Whilton Production: Irene Herlihy ‘Manufacturing: Lisa Flanagan ‘Typeset by Best-set Typesetter Ltd., Hong Kong. Printed and bound by Maple-Vail Printed in the United States of America 00 01 02 5432 ‘The Blackwell Science logo is a trade mark of Blackwell Science Ltd., registered at the United Kingdom Trade Marks Registry Library of Congress Cataloging-in-Publication Data Veterinary microbiology / edited by Dwight C. Hirsh, Yuan Chung Zee. Pp. cm. Includes bibliographical references and index. ISBN 0-86542-543-4 1, Veterinary microbiology. I. Hirsh, Dwight C. IL. Zee, ¥. C. SE780.2.V48 1999 636.089°601 — de21 98-30284 cp Notice: The indications and dosages of all drugs in this book have been recommended in the medical literature and conform to the practices of the general community. The medications described do not necessarily have specific approval by the Food and Drug Administration for use in the diseases and dosages for which they are rec- ‘ommended. The package insert for each drug should be consulted for use and dosage as approved by the FDA. Because standards for usage change, itis advisable to keep abreast of revised recommendations, particularly those concerning new drugs. ‘To Lucy, Dwight, and Elizabeth for years of patience and understanding, and To Elizabeth, Norman, and Charlie for instilling a love for all creatures Contents Contributors Preface PARTI INTRODUCTION Chapter 1 Parasitism and Pathogenicity ERNST L. BIBERSTEIN Chapter 2 Immune Responses to Infectious Agents Lauri. J. GERSHWIN Chapter 3 Laboratory Diagnosis Dwi C. Hirsi, YUAN CHUNG ZEE, AND ANTHONY E. CASTRO Chapter 4 Antimicrobial Chemotherapy JouN B. Prescorr Chapter § Antimicrobial Drugs: A Strategy for Rational Use and the Ramifications of Misuse DwiGHT C. Hinst Chapter 6 Vaccines YUAN CHUNG Zee AND Dwicitt C. HiRst PART Il BACTERIA AND FUNGI Chapter 7 The Alimentary Canal as a Microbial Habitat DwicHt C. Hirst Chapter 8 Family Enterobacteriacene Dwrcnt C. Hirst Chapter 9 Escherichia DwiGHT C. Hinstt Chapter 10 Salmonella wich C, Hinsit Chapter 11. Shigella wich C, Hist Chapter 12 Non-Spore-Forming Obligate Anaerobes of the Alimentary Tract Dwicht C. Hist Chapter 13 Serpulina Dwicit C. Hirst Chapter 14 Spiral Organisms I: Campylobacter = Arcobacter — Lawsonia (Digestive Tract) Dwicttr C. Hirst Chapter 15 Spiral Organisms Il: Helicobacter James G. Fox Chapter 16 Pseudomonas Dwicet C. Hiast 1s 28, 46 st 59 6 65 69 78 83 86 89. 93 100 Chapter 17 Yersinia enterocolitica Dwicst C. Hirst Chapter 18 Mycobacterium avium ssp. paratuberculosis (Mycobacterium pparatuberculosis) Dwicht C. Hirst Chapter 19 Candida ERNST L. BIBERSTEIN Chapter 20 The Respiratory Tract as a Microbial Habitat BANST L. BIBERSTEIN Chapter 21 Staphylococei Exyst L. BIBERSTEIN AND DwiGHT C. HiRst Chapter 22 Streptococci Exnst L. BIneRSTEIN AND DwiGHt C. HiRst Chapter 23 Corynebacteria; Arcanobacterium (Actinomyces) pyogenes; Rhodococcus equi [EnxsT L. BIgERSTEIN AND DwiGHt C. Hinsti Chapter 24 Pasteurella Exnst L. BineRSTEIN AND Dwicht C. HiRst Chapter 25 Actinobacillus Ennst L. BIBERSTEIN AND DWIGHT C. HIRSH Chapter 26 Haemophilus spp. Enxst L. BIBERSTEIN Chapter 27 Bordetella Enxst L. BIBERSTEIN AND DWIGHT C. Hinstt Chapter 28 Moraxella Exnst L. BIBERSTEIN AND DWIGHT C. HiRsHt Chapter 29 Burkholderia mallei and Burkholderia pseudomallei ERNST L, BIBERSTEIN AND DWIGHT C. HIRst ‘Chapter 30 Mycobacterium Species: The Agents of Animal Tuberculosis Exnst L, BIBERSTEIN AND DWIGHT C. HIRSH Chapter 31 Mollicutes icHaRD L. WALKER Chapter 32 Chlamydiae Exwst L. BIBERSTEIN AND DwiGHT C. HiRstt Chapter 33 The Urinary Tract as a Microbial [ERNST L. BIBERSTEIN AND DWIGHT C. HIRSH Chapter 34 Leptospirae Rance B, LEFEBVRE. Chapter 35 ‘The Genital Tract as a Microbial Habitat Dwi C. Hirst 102 104 109 3 us 120 127 135 ut 144 148 151 155 158 165 173 178 185 190 vill Contents Chapter 36 Campylobacter — Arcobacter (Reproductive Tract) Dwi C. Hirst Chapter 37 Brucella RICHARD L. WALKER Chapter 38 Taylorella equigenitalis ERNsT L. BIBERSTEIN Chapter 39 The Skin as a Microbial Habitat: Bacterial Skin Infections ERNST L.. BIBERSTEIN| Chapter 40 Dermatophytes ERNST L. BIBERSTEIN Chapter 41 Agents of Subcutaneous Mycoses [ERNST L. BIBERSTE! Chapter 42 Listeria RICHARD L. WALKER Chapter 43 Erysipelothrix RICHARD L. WALKER Chapter 44 The Clostridia ERNST L. BIBERSTEIN AND DWIGHT C. HIRSH Chapter 45 The Genus Bacillus ERNST L. BIBERSTEIN AND DWIGHT C. HiRsHt Chapter 46 Pathogenic Actinomycetes (Actinomyces and Nocardia) ERNST L, BIBERSTEIN AND DWIGHT C. Hirsit Chapter 47 Agents of Systemic Mycoses ERNST L., BIBERSTEIN Chapter 48 Mycotoxins Francis D. GaLey Chapter 49 The Yersiniae [ERNST L. BISERSTEIN AND DWIGHT C. HIRSH Chapter 50 Francisella tularensis [ERNST L. BIBERSTEIN Chapter $1 Borrelia spp. RANE B. LEFEBVRE Chapter 52 Streptobacillus moniliformis [ERNST L. BISERSTEIN Chapter 53 Rickettsial Agents of Animal Disease; the Rickettsieae [ERNST L. BIBERSTEIN AND DWIGHT C. HIRSH Chapter $4 Ehrlichieae: Ehrlichia, Cowaria, and Neorickettsia ERNST L. BIBERSTEIN AND DWIGHT C. HIRSH Chapter SS Bartonellaceae BRUNO B. CHOMEL Chapter 56 Anaplasmataceae ERNST L. BIBERSTEIN 192 196 204 206 214 220 228 229 233, 246 250 256 274 281 285 287 290 291 204 299 304 PART Ill VIRUSES Chapter 57 General Properties of Viruses JANET S. BUTEL, JOSEPH L. MELNICK, AND YUAN CHUNG ZEE Chapter 58 Pathogenesis of Viral Diseases ‘YUAN CHUNG ZEE Chapter 59 Parvoviridae YuAN CHUNG Zee Chapter 60 Iridoviridae Jersaey L. Storr Chapter 61 Papovaviridae ‘YUAN CHUNG Zee Chapter 62 Adenoviridae YUAN CHUNG ZeE Chapter 63 Herpesviridae ALEX A. ARDANS Chapter 64 Poxviridae Jereeey L. Storr Chapter 65 Picornaviridae Jererey L. Storr Chapter 66 Caliciviridae YUAN CHUNG Zee Chapter 67 Togaviridae and Flaviviridae Jesrary L. STOTT Chapter 68 Orthomyxoviridae ALEX A. ARDANS. Chapter 69 Paramyxoviridae YUAN CHUNG Zee Chapter 70 Rhabdoviridae Yuan CHUNG ZEE Chapter 71 Coronaviridae Jerrrey L, Storr Chapter 72 Reoviridae Jesrney L. Storr Chapter 73 Biraviridae Jesrney L. Storr Chapter 74 Retroviridae RicHarD M. Donovan Chapter 75 ‘Transmissible Spongiform Encephalopathies ‘YUAN CHUNG Zee Index 309 ai 328 333 340 as 346 350 365 an 379 385 396 403 a2 ans 430 439 492 461 463 Contributors Alex A. Ardans, DVM, MS Professor, Department of Medicine and Epidemiology Director, California Veterinary Diagnostic Laboratory System, School of Veterinary Medicine University of California Davis, California Emnst L. Biberstein, DVM, PhD Professor Emeritus, Department of Pathology, Microbiology, and Immunology School of Veterinary Medicine University of California Davis, California Janet S. Butel, PhD Professor of Virology Division of Molecular Virology Baylor College of Medicine Houston, Texas Anthony E. Castro, DVM, PhD Department of Veterinary Sciences, Pennsylvania State University University Park, Pennsylvania Bruno B. Chomel, DVM, PhD Associate Professor, Department of Population Health and Reproduction, ‘School of Veterinary Medicine University of California Davis, California Richard M. Donovan, PhD Director of Infectious Diseases Research Henry Ford Health Sciences Center Detroit, Michigan. James G. Fox, DVM Professor and Director, Division of Comparative Medicine Massachusetts Institute of Technology Cambridge, Massachusetts Francis D. Galey, DVM, PhD Associate Professor, Department of Molecular Biosciences Section Head, Clinical Toxicology California Veterinary Diagnostic Laboratory School of Veterinary Medicine University of California Davis, California Laurel J. Gershwin, DVM, PhD Professor, Department of Pathology, Microbiology, and Immunology Chief of Service, Clinical Immunology and Virology Laboratory Veterinary Medical Teaching Hospital ‘School of Veterinary Medicine University of California Davis, California Dwight C. Hirsh, DVM, PhD Professor, Department of Pathology, Microbiology, and Immunology Chief of Service, Clinical Microbiology Laboratory ‘Veterinary Medical Teaching Hospital ‘School of Veterinary Medicine University of California Davis, California Rance B. LeFebvre, PhD Professor, Department of Pathology, Microbiology, and Immunology ‘School of Veterinary Medicine University of California Davis, California Joseph L. Melnick, PhD Professor of Virology and Epidemiology Baylor College of Medicine Houston, Texas John F. Prescott, Vet MB, PhD Professor, Department of Pathobiology University of Guelph Ontario, Canada Jeffrey L. Stott, PhD Professor, Department of Pathology, Microbiology, and Immunology School of Veterinary Medicine University of California Davis, California Richard L. Walker, DVM, PhD, MPVM Associate Professor Department of Pathology, Microbiology, and Immunology. School of Veterinary Medicine; Section Head, Bacteriology California Veterinary Diagnostic Laboratory University of California Davis, California ‘Yuan Chung Zee, DVM, PhD Professor, Department of Pathology, Microbiology, and Immunology School of Veterinary Medicine University of California Davis, California Preface ‘This book is intended primarily for veterinary students, to accompany and supplement their first courses in pathogenic bacteriology-mycology and virology. Its focus includes pathogenic mechanisms and processes in infec- tious diseases; methods of diagnosis; and principles of resistance, prevention, and therapy. A working knowl edge of general microbiology is assumed. Beyond serving as a resource for students, the book is also meant to serve as a convenient reference for veteri- narians and veterinary scientists whose main line of activ- ity and expertise is outside the areas of microbiology. ‘The manner of presentation, i.e, sequence and chapter organization, was determined by the way which the editors found most appropriate for teaching their respec- tive subjects: the bacteriology-mycology portion is arranged roughly by host organ systems and other milieus that serve as sources of pathogenic agents. This approach creates a logical place to consider the various environ- ‘ments as microbial habitats. The virology section is orga- nized more along taxonomic lines. Regardless of the user's preference or custom, all topics are readily located ‘with the aid of the table of contents and the alphabetic index. ‘While we have included all agents likely to be encoun- tered in veterinary practice, we have tried to avoid indis- criminate listing of conditions and microorganisms reported only exceptionally, particularly in the contem- porary literature. Our objective is the main current of veterinary microbiology. ‘The purpose of the reference citations at the end of chapters is to guide the reader to more comprehensive sources of information rather than to document chapter content. We have therefore favored recent reviews and monographs. We believe these to be of greatest use since they will lead the interested reader to the primary sources, which considerations of brevity and economy forced us to omit. The content of this book varies somewhat from our ‘earlier work, Review of Veterinary Microbiology (1990). Most notable is the replacement of the chapters dealing with ‘general immunology with one limited to a discussion of immunologic phenomena related to infectious agents. We have also changed the focus of the chapters dealing ‘with antimicrobial agents to one more clinical by the addition of a chapter on the rational choice of antimi- ‘crobial agents in the treatment of an infectious disease. ‘We have added chapters dealing with microbiological diagnosis, and one on vaccines. ‘We gratefully acknowledge Trudi Schuster, whose help Js much appreciated. A special thank you goes to Jill Connor and Irene Herlihy of Blackwell Science, who have been unbelievably patient and extremely helpful in getting our effort to press. DCH. ¥.CL. PART I Introduction 1 Parasitism and Pathogenicity ERNST L. BIBERSTEIN Veterinary microbiology deals with microbial agents affecting animals. Such agents are characterized accord- ing to their ecologic arrangements: parasites live in per- manent association with, and at the expense of, animal hosts; saprophytes normally inhabit the inanimate envi- ronment. Parasites that cause their host no discernible harm are called commensals. The term symbiosis usually refers to reciprocally beneficial associations of organisms. This arrangement is also called rmutualism. Pathogenic organisms are parasites or saprophytes that ‘cause disease. The process by which they establish them- selves in a host individual is infection, but infection need not be followed by clinical illness. The term virulence is sometimes used to mean pathogenicity but sometimes to express degrees of pathogenicity. SOME ATTRIBUTES OF HOST—PARASITE RELATIONSHIPS Many pathogenic microorganisms are host-specific in that they parasitize only one or a few animal species. Streptococcus equt is essentially limited to horses. Others — certain Salmonella types, for example — have a broad host range. The basis for this difference in host specificity is incompletely understood, but it may in part be related to the need for specific attachment devices between host (receptors) and parasite (adhesins). Some agents infect several host species but with varying effects. The plague bacillus Yersinia pestis behaves as a commensal parasite in many, but by no means all, small rodent species but causes fatal disease in rats and humans. Evolutionary pressure may have produced some Of these differences, but not others: Coccidioides immitis, a saprophytic fungus requiring no living host, infects cattle and dogs with equal ease; yet it produces no clini- cal signs in cattle but frequently causes progressive fatal disease in dogs. Potential pathogens vary in their effects on different tissues in the same host. The Escherichia coli that is com- ‘mensal in the intestine can cause severe disease in the ary tract and peritoneal cavity. ‘Some microorganisms that are commensal in one habitat may turn pathogenic in a habitat that is patho- logically altered or otherwise compromised. Thus, oral streptococci, which occasionally enter the bloodstream, may colonize a damaged heart valve and initiate bacte- rial endocarditis. In the absence of such a lesion, however, ‘they would be cleared uneventfully via the macrophage system. Similarly, the frequent entrance of intestinal bac- teria into vascular channels normally leads to their dis- posal by humoral and cellular defense mechanisms. In immuno-incompetent hosts, however, such entrance may lead to fatal septicemia, ‘Transfer to a new host of tissue, or a change in host resistance, are common ways that commensal parasites are converted into active pathogens. Commensalism is the stable form of parasitic existence. It ensures survival of, the microorganism, which active disease would jeopar- dize by killing the host or evoking an active immune response. Either effect deprives the agent of its habitat. Evolutionary selective pressure therefore tends to elimi- nate host-parasite relationships that threaten the survival of either partner. It does so by allowing milder strains of the pathogen, which permit longer survival of the host and thereby facilitate their own dissemination, to replace the more lethal ones. It also favors a resistant host population by screening out highly susceptible stock. ‘The trend is thus toward commensalism. Most agents causing serious infections have alternative modes of sur- vival as commensals in tissues or hosts not subject to disease (e-g., plague) or in the inanimate environment (eg,, coccidioidomycosis). Others cause chronic infec- tions lasting months or years (tuberculosis, syphilis), during which their dissemination to other hosts ensures their survival. CRITERIA OF PATHOGENICITY — KocH’s PosTULaTEs ‘The presence of a microorganism in diseased individuals does not prove its pathogenic significance. To demon- strate the causal role of an agent in a disease, the follow- ing qualifications or “postulates” formulated by Robert Koch (1843-1910) should be satisfied: 1. The suspected agent is present in all cases of the disease. 2. The agent is isolated from such disease and propagated serially in pure culture, apart from its natural host. 3. Upon introduction into an experimental host, the isolate produces the original disease. 4, The agent can be reisolated from this experimental infection. 4 Part I Introduction ‘These postulates are ideals that are not satisfied in all cases of infectious diseases. The presence of some microorganisms cannot be demonstrated at the time of disease, especially in affected tissues (tetanus, botulism). Others lose virulence rapidly after isolation (Leptospira spp), while still others, though indispensable for pathogenesis, require undetermined accessory factors (Pasteurella-related pneumonias), For some human viral pathogens (cytomegalovirus), no experimental host is known, and some agents (e.., Mycobacterium leprae) have not been grown apart from their natural hosts. ELEMENTS IN THE PRODUCTION OF AN INFECTIOUS DISEASE Effective transmission through indirect contact occurs by ingestion; inhalation; or mucosal, cutaneous, or wound contamination. Airborne infection takes place largely via

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