Veterinary MicrobiologyVETERINARY 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 creaturesContents
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
190vill 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
463Contributors
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, CaliforniaPreface
‘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
Introduction1 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