FOODBORNE PARASITES

FOOD MICROBIOLOGY AND FOOD SAFETY SERIES

Food Microbiology and Food Safety publishes valuable, practical, and timely
resources for professionals and researchers working on microbiological
topics associated with foods, as well as food safety issues and problems.

Series Editor

Michael P. Doyle, Regents Professor and Director of the Center for Food Safety,
University of Georgia, Griffin, GA, USA

Editorial Board

Francis F. Busta, Director – National Center for Food Protection and Defense,
University of Minnesota, Minneapolis, MN, USA
Bruce R. Cords, Vice President, Environment, Food Safety & Public Health,
Ecolab Inc., St. Paul, MN, USA
Catherine W. Donnelly, Professor of Nutrition and Food Science, University
of Vermont, Burlington, VT, USA
Paul A. Hall, Senior Director Microbiology & Food Safety, Kraft Foods North
America, Glenview, IL, USA
Ailsa D. Hocking, Chief Research Scientist, CSIRO—Food Science Australia,
North Ryde, Australia
Thomas J. Montville, Professor of Food Microbiology, Rutgers University,
New Brunswick, NJ, USA
R. Bruce Tompkin, Formerly Vice President-Product Safety, ConAgra Refrig-
erated Prepared Foods, Downers Grove, IL, USA

Titles

PCR Methods in Foods, John Maurer (Ed.) (2006)

Viruses in Foods, Sagar M. Goyal (Ed.) (2006)
Foodborne Parasites, Ynes R. Ortega (Ed.) (2006)
FOODBORNE PARASITES

Edited by
Ynes R. Ortega
University of Georgia, Griffin, GA, USA
Ynes R. Ortega
Center for Food Safety
Department of Food Science and Technology
University of Georgia
1109 Experiment St.
Griffin, GA 30223
Ortega@uga.edu

Library of Congress Control Number: 2006921159

ISBN-10: 0-387-30068-6 e-ISBN 0-387-31197-1 Printed on acid-free paper.

ISBN-13: 978-0387-30068-9

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 Preface

Microbiologists are being challenged as foodborne outbreaks are increasingly be-

ing observed worldwide. Most of these outbreaks are associated with viral and
bacterial pathogens such as Campylobacter, Salmonella, and lately Escherichia coli
O157:H7, which emerged in the 1990s. The role of food in transmission of parasites
was not studied until later.
Although parasites have been evolving with man since antiquity, the control
and eradication of these diseases is still far from being achieved and they are more
frequently being reported in the literature as causative agents of food and waterborne
illnesses.
Parasites have been consistently reported in developing and endemic countries.
However, the presence of these parasites in the developed world has increased in
the past two decades. For example, in the United States alone, foodborne outbreaks
have multiplied by more than a factor of eight during the past 15 years.
Many factors can be attributed to the rise in parasitic foodborne outbreaks. The
increase of international travel and population migration encourages rapid disease
dissemination. Globalization of the food supply has introduced new challenges in
the U.S. food safety arena. The food habits of Americans are changing, resulting
in fresh foods and vegetables being consumed. Consequently, importation of foods
is now necessary in order to satisfy the consumer demands for a year-round sup-
ply of certain commodities, such as exotic fruits and vegetables. Importations have
also increased as the costs of particular products have been reduced. Unfortunately,
transportation conditions such as controlled refrigeration favor survival of parasites
on fruits and vegetables. In addition, restaurants presenting ethnic foods not tradi-
tionally consumed are revealing public health issues that have not been considered.
For example, Sarcocystis, a nematode parasite, has been identified in meats served
at Arab restaurants in Brazil where meats are consumed raw.
Moreover, populations are at increased risk for acquiring infection and develop-
ing gastrointestinal illness due to the prevalence of asymptomatic carriers and the
zoonotic potential of some parasites that contribute to the spread of the pathogen. For
example, inadequate composting and lack of agricultural practices has contributed
to the exposure of food products to animal waste and hence, parasites.
Medical advances have contributed to changes in the US demographics, yet, the
elderly, immunocompromised individuals and children are susceptible populations
to parasitic infections.
Misdiagnosis of parasitic infections is not uncommon. Diagnostic laboratories
and medical personnel may not be familiar with parasite identification and the risk
factors for acquiring these infections. Excretion of parasites is intermittent, requiring
that more than one stool sample be examined. In some instances, special staining
procedures not routinely requested by medical personnel are required to identify
parasites properly.

v
vi P r e fac e

Outbreak investigations may take many days, by which time samples of the im-
plicated product may not be available for investigation. Therefore, food scientists
and the medical community need to be aware of the role of parasites as significant
agents of foodborne outbreaks. Routine examination of clinical specimens of indi-
viduals with gastrointestinal illness does not search for parasites. In fact, testing for
Cyclospora and Cryptosporidium must be specifically requested. Improvements in
detection systems not only in the clinical settings, but also in the food microbiol-
ogy laboratories have allowed for identification of infections that previously went
undetected.
Molecular assays have been very helpful in foodborne outbreak investigations
but are limited by isolation procedures for the parasites. Recovery is particularly
important since parasites are generally inert in the environment, and enrichment
procedures applied to bacterial contaminants are not an option.
In the 1980s, Cryptosporidium emerged as a significant pathogen responsible for
high mortality in immunocompromised patients; particularly in those with AIDS.
Although to date there is no effective therapy for cryptosporidiosis, a great deal
of information was learned about this parasite, most notably that the cellular im-
mune response plays a significant role in eradication. Cyclospora, another emerging
parasite, has also proven to be a challenge to work with and has demonstrated to
be highly resistant to environmental conditions. Cyclospora is a parasite that is
associated with foodborne outbreaks in the United States and is caused by con-
taminated produce imported for human consumption. Trichinella, a nematode, is
another emerging pathogen that is frequently reported in the United States and is
commonly associated with the ingestion of game meats and sausages that are not
properly cooked.
This book will review the two major parasite groups that are transmitted via water
or foods: the protozoa, which are single celled organisms and the helminths. The
helminths are classified into three sub groups, the cestodes (tapeworms), nematodes
(round worms), and trematodes (flukes). To better understand their significance, each
chapter will be covering the biology, mechanisms of pathogenesis, epidemiology,
treatment, and inactivation of these parasites. A better understanding of the biology
and control of parasitic infections is necessary to reduce and eliminate outbreaks in
the United States and elsewhere.

PROTOZOA
Protozoa are single-celled organisms. They are eukaryotes which have compartmen-
talized organelles and the infectious stages are environmentally resistant. Protozoa
can be blood borne or transmitted via a vector, such as a mosquito. The most well-
known disease in this group is malaria. Other protozoa, acquired via water, are
environmentally resistant, and can be responsible for worldwide human illness. A
classic example of this group is Cryptosporidium. As we learn more of the molecular
composition of these parasites, and the development of molecular tools for geno-
typing, new species are being described. Along with these, a reevaluation of these
species and their significance in public health is changing. This is particularly true
 P r e fac e vii

in the case of Cryptosporidium. The species C. parvum was considered to be unique,

although there was significant evidence that the clinical presentation was variable
as well as host susceptible. Now more genotypes are being identified in human and
animal species. Although most of the outbreaks associated with Cryptosporidium
have been waterborne, a few reports have been foodborne. The contamination has
been associated with food preparation practices, hygienic conditions of the food
handlers, or by fresh produce that may have been contaminated on the farms.
Most parasites are obligate intracellular organisms. In contrast with bacteria,
parasites are inert and do not multiply in the environment. Therefore, studies of
food matrices and determination of the safety of some products is challenging.
Isolation and detection procedures are crucial because an enrichment process for
parasites is not available. Although molecular assays overcome these difficulties,
specific limitations of each organism will be discussed in the pertinent chapters.
Other protozoa can be acquired by more than one route. Toxoplasma is a good
example of this group. Present in almost every region of the world, Toxoplasma
can be acquired by most warm-blooded animals, by ingestion of oocyst (the en-
vironmentally resistant form) in water or fresh produce, or by ingestion of raw or
inadequately cooked meats which contain tissue cysts of the parasite.
Most of the foodborne protozoa cause diarrheal illness in humans; however,
Toxoplasma can have detrimental consequences for the fetus of pregnant women
and in immunocompromised individuals. In immunocompetent individuals toxo-
plasmosis may be asymptomatic, but if the immune competency diminishes either
by chemotherapy or by acquired deficiency, the individual can present symptomatic
toxoplasmosis that is most commonly manifested as encephalitis.

HELMINTHS
This group is commonly identified as worms and is rapidly emerging in the United
States and worldwide. Ingestion of raw or improperly cooked meats may be the
source of infection with most of these parasites.
Helminths have very complex life cycles and in most instances humans are
accidental hosts. However, in other cases the adult worms can find a home in the
human body, start reproducing and shed eggs in the environment. These eggs could
infect other susceptible individuals.
Adult nematodes, present as male and female worms, can be found in the gut of
the infected individual. In contrast, cestodes and trematodes can be hermaphrodites
and self fertilize their eggs; therefore, most of them do not require a male and female
worm to complete their life cycle.
Some infections with helminths have been associated with immigrant popula-
tions that initiate an outbreak or start developing clinical signs infrequently observed
by medical personnel. An example of this is cysticercosis. Outbreaks in the United
States were described in an ethnic group that did not consume pork, and further
investigation showed that infection was initiated by a foreign housekeeper who had
teniasis.
viii P r e fac e

Trematoda can also be acquired by ingestion of raw or undercooked fish. Al-

though freezing can inactivate the cysts, the larval stages can remain viable at re-
frigeration temperatures. Many consumers prefer fresh fish due to the detrimental
textural changes in the fish after freezing.

Ynes R. Ortega
 Contributors

Ann M. Adams, US Food and Drug Administration, Kansas City District, Lenexa,
KS 66214
Natalie Bowman, Columbia College of Physicians and Surgeons, Columbia Uni-
versity, New York, NY 10032
Vitaliano Cama, Department of International Health, Bloomberg School of Public
Health, Johns Hopkins University, Baltimore, MD 21205
George D. Di Giovanni, Agricultural Research and Extension Center, Texas A&M
University, El Paso, TX 79927
Joseph Donroe, Tufts University School of Medicine, Tufts University School of
Public Health, Boston, MA 02111
Robert Gilman, Department of International Health, Bloomberg School of Public
Health, Johns Hopkins University, Baltimore, MD 21205
Kristina D. Mena, School of Public Health, University of Texas Health Science
Center, El Paso, Texas 79902
Ynes R. Ortega, Center for Food Safety, Department of Food Science and Tech-
nology, University of Georgia, Griffin, GA 30223
Charles R. Sterling, Department of Veterinary Science and Microbiology, Univer-
sity of Arizona, Tucson, AZ 85721
Gregory D. Sturbaum, CH Diagnostic and Consulting Service, Inc. Loveland, CO
80537
Irshad M. Sulaiman, Biotechnology Core Facility Branch, Scientific Resources
Program, National Center for Infectious Diseases, Centers for Disease Control and
Prevention, Atlanta, GA 30333
Lihua Xiao, Division of Parasitic Diseases, Centers for Disease Control and Pre-
vention, Atlanta, GA 30341

ix
 Contents

Chapter 1. Amoeba and Ciliates

Ynes R. Ortega . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2 Amoeba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2.1 Entamoeba histolytica . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.3 Dientamoeba fragilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1.3.1 Morphology and Transmission . . . . . . . . . . . . . . . . . . 6
1.3.2 Therapy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.4 Nonpathogenic Amoeba . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.4.1 Entamoeba hartmanni . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.4.2 Entamoeba coli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.4.3 Endolimax nana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.4.4 Iodamoeba butschlii . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.5 Free-Living Amoebae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1.6 Ciliates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1.6.1 Life Cycle and Morphology . . . . . . . . . . . . . . . . . . . . 9
1.6.2 Clinical Significance . . . . . . . . . . . . . . . . . . . . . . . . . . 9
1.6.3 Diagnosis and Treatment . . . . . . . . . . . . . . . . . . . . . . . 11
1.6.4 Epidemiology and Prevention . . . . . . . . . . . . . . . . . . . 11
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Chapter 2. The Biology of Giardia Parasites

Irshad M. Sulaiman and Vitaliano Cama . . . . . . . . . . . . . . . . . . . . 15
2.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
2.2 Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
2.3 Detection and Classification of Giardia . . . . . . . . . . . . . . . . 17
2.3.1 Detection Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
2.3.2 Classification of G. intestinalis . . . . . . . . . . . . . . . . . . 19
2.3.3 Genotyping of G. intestinalis . . . . . . . . . . . . . . . . . . . 20
2.4 Transmission and Epidemiology . . . . . . . . . . . . . . . . . . . . . . 23
2.4.1 Human . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
2.4.2 Environmental . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
2.5 Control and Treatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

Chapter 3. Coccidian Parasites

Vitaliano Cama . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
3.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
3.2 Background/History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

xi
xii Contents

3.3 Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
3.4 Clinical Significance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
3.5 Transmission and Epidemiology . . . . . . . . . . . . . . . . . . . . . . 39
3.5.1 Cyclospora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
3.5.2 Isospora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
3.5.3 Sarcocystis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
3.6 Diagnosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
3.7 Treatment and Control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

Chapter 4. Cryptosporidium and Cryptosporidiosis

Lihua Xiao and Vitaliano Cama . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
4.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
4.2 Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
4.3 Life Cycle and Developmental Biology . . . . . . . . . . . . . . . . 60
4.4 Epidemiology and Transmission . . . . . . . . . . . . . . . . . . . . . . 61
4.4.1 Cryptosporidiosis in Immunocompetent Persons . . . . 61
4.4.2 Cryptosporidiosis in Immunocompromised
Persons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
4.4.3 Transmission Routes and Infection Sources:
Anthroponotic Versus Zoonotic Transmission . . . . . . 63
4.4.4 Waterborne Transmission . . . . . . . . . . . . . . . . . . . . . . 64
4.4.5 Foodborne Transmission . . . . . . . . . . . . . . . . . . . . . . . 66
4.5 Detection and Diagnosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
4.5.1 Serologic Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
4.5.2 Methods for Detection of Cryptosporidium
in Stool Specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
4.5.3 Methods for Detection of Cryptosporidium Oocysts
in Environmental Samples . . . . . . . . . . . . . . . . . . . . . . 77
4.6 Treatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
4.7 Control of Cryptosporidium Contamination in
Water and Food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

Chapter 5. Toxoplasmosis
Ynes R. Ortega . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
5.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
5.2 Parasite Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
5.3 Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
5.4 Transmission . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
5.5 Identification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
5.5.1 Molecular Assays . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
5.5.2 Riboprinting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
5.6 Pathogenicity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
 Contents xiii

5.7 Epidemiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115

5.7.1 Humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
5.7.2 Swine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
5.7.3 Poultry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
5.7.4 Sheep and Goats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
5.7.5 Other Animal Species . . . . . . . . . . . . . . . . . . . . . . . . . 122
5.8 Treatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123
5.9 Inactivation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

Chapter 6. Food-Borne Nematode Infections

Charles R. Sterling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
6.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
6.2 Trichinella spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
6.2.1 Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
6.2.2 Speciation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
6.2.3 Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
6.2.4 Epidemiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
6.2.5 Human Trichinellosis–Epidemiology . . . . . . . . . . . . . 140
6.2.6 Clinical Manifestations . . . . . . . . . . . . . . . . . . . . . . . . 142
6.2.7 Diagnosis and Treatment . . . . . . . . . . . . . . . . . . . . . . . 143
6.2.8 Prevention and Control . . . . . . . . . . . . . . . . . . . . . . . . 143
6.3 Anisakis simplex and
Related Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
6.3.1 Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
6.3.2 Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
6.3.3 Epidemiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
6.3.4 Clinical Manifestations . . . . . . . . . . . . . . . . . . . . . . . . 147
6.3.5 Diagnosis and Treatment . . . . . . . . . . . . . . . . . . . . . . . 147
6.3.6 Prevention and Control . . . . . . . . . . . . . . . . . . . . . . . . 148
6.4 Angiostrongylus cantonensis and
Angiostrongylus costaricensis . . . . . . . . . . . . . . . . . . . . . . . . 148
6.5 Gnathostoma spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
6.6 Gongylonema spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
6.7 Other Nematode Infections with Food-Borne
Associations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153

Chapter 7. Foodborne Trematodes

Ann M. Adams . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
7.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
7.2 Paragonimus spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
7.2.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
7.2.2 Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
xiv Contents

7.2.3 Epidemiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165

7.2.4 Clinical Signs, Diagnosis, and Treatment . . . . . . . . . . 166
7.3 Clonorchis sinensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
7.3.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
7.3.2 Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
7.3.3 Epidemiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170
7.3.4 Clinical Signs, Diagnosis, and Treatment . . . . . . . . . . 171
7.4 Opisthorchis viverrini and Opisthorchis felineus . . . . . . . . . 174
7.4.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174
7.4.2 Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174
7.4.3 Epidemiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
7.4.4 Clinical Signs, Diagnosis, and Treatment . . . . . . . . . . 176
7.5 Nanophyetus salmincola . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
7.5.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
7.5.2 Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
7.5.3 Salmon Poisoning Disease . . . . . . . . . . . . . . . . . . . . . 179
7.5.4 Clinical Signs, Diagnosis, and Treatment . . . . . . . . . . 180
7.6 Fasciola spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
7.6.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
7.6.2 Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
7.6.3 Epidemiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
7.6.4 Clinical Signs, Diagnosis, and Treatment . . . . . . . . . . 184
7.7 Fasciolopsis buski . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186
7.7.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186
7.7.2 Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
7.7.3 Epidemiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
7.7.4 Clinical Signs, Diagnosis, and Treatment . . . . . . . . . . 188
7.8 Prevention and Control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191

Chapter 8. Cestodes
Natalie Bowman, Joseph Donroe, and Robert Gilman . . . . . . . . . . 197
8.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
8.2 Taenia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
8.2.1 Taenia solium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
8.2.2 Taenia saginata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
8.2.3 Taenia asiatica. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
8.3 Diphyllobothrium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
8.3.1 Diphyllobothrium latum. . . . . . . . . . . . . . . . . . . . . . . . 211
8.3.2 Diphyllobothrium pacificum. . . . . . . . . . . . . . . . . . . . . 214
8.4 Spirometra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
8.4.1 Spirometra mansoides. . . . . . . . . . . . . . . . . . . . . . . . . 215
8.5 Echinococcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
8.5.1 Echinococcus granulosus. . . . . . . . . . . . . . . . . . . . . . . 216
8.5.2 Echinococcus multilocularis. . . . . . . . . . . . . . . . . . . . 221
 Contents xv

8.6 Hymenolepis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223

8.6.1 Hymenolepis nana. . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225

Chapter 9. Waterborne Parasites and Diagnostic Tools

Gregory D. Sturbaum and George D. Di Giovanni . . . . . . . . . . . 231
9.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
9.2 Parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
9.3 Concentration and Isolation Techniques . . . . . . . . . . . . . . . 235
9.4 Detection Methodologies . . . . . . . . . . . . . . . . . . . . . . . . . . 237
9.4.1 Microscopic Techniques . . . . . . . . . . . . . . . . . . . . . . 237
9.4.2 Nucleic Acid Techniques . . . . . . . . . . . . . . . . . . . . . . 239
9.4.3 Immunological-based Techniques . . . . . . . . . . . . . . . 248
9.4.4 Viability Techniques . . . . . . . . . . . . . . . . . . . . . . . . . 250
9.5 Proper Evaluation (QA/QC) . . . . . . . . . . . . . . . . . . . . . . . . 253
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255

Chapter 10. Risk Assessment of Parasites in Food

Kristina D. Mena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
10.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
10.2 The Risk Assessment Framework . . . . . . . . . . . . . . . . . . . . 276
10.2.1 Defining the Hazard . . . . . . . . . . . . . . . . . . . . . . . . . 276
10.2.2 Exposure Assessment . . . . . . . . . . . . . . . . . . . . . . . 277
10.2.3 Hazard Characterization (Dose-response
Assessment) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
10.2.4 Risk Characterization . . . . . . . . . . . . . . . . . . . . . . . 280
10.2.5 Assumptions, Assumptions, Assumptions . . . . . . . 281
10.2.6 Emerging Applications of Microbial Risk
Assessment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285