Cryptosporidium and Cryptosporidiosis 2nd Edition
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CRYPTOSPORIDIUM
AND
CRYPTOSPORIDIOSIS
Second Edition
Edited by
Ronald Fayer
Lihua Xiao
Boca Raton London New York
CRC Press is an imprint of the
Taylor & Francis Group, an informa business
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CRC Press
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© 2008 by Taylor & Francis Group, LLC
CRC Press is an imprint of Taylor & Francis Group, an Informa business
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International Standard Book Number-13: 978-1-4200-5226-8 (Hardcover)
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Library of Congress Cataloging-in-Publication Data
Cryptosporidium and cryptosporidiosis / [edited by] Ronald Fayer, Lihua Xiao. -- 2nd ed.
p. ; cm.
“A CRC title.”
Includes bibliographical references and index.
ISBN 978-1-4200-5226-8 (hardcover : alk. paper)
1. Cryptosporidiosis. 2. Cryptosporidium. I. Fayer, R. II. Xiao, Lihua, 1962- III. Title.
[DNLM: 1. Cryptosporidiosis. 2. Cryptosporidium. WC 730 C956 2007]
RC136.5.C79 2007
616.9’36--dc22 2007021158
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Ernest Edward Tyzzer
(Courtesy of the Francis A. Countway Library of Medicine, Boston, Massachusetts)
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Contents
1 General Biology ....................................................................................................1
Ronald Fayer
2 Genomics .............................................................................................................43
Jessica C. Kissinger
3 Biochemistry .......................................................................................................57
Guan Zhu
4 Epidemiology ......................................................................................................79
Gordon Nichols
5 Molecular Epidemiology ................................................................................. 119
Lihua Xiao and Una M. Ryan
6 Diagnostics ........................................................................................................ 173
Huw Smith
7 Immune Responses .......................................................................................... 209
Vincent McDonald
8 Clinical Disease and Pathology...................................................................... 235
Cirle Alcantara Warren and Richard L. Guerrant
9 Prophylaxis and Chemotherapy...................................................................... 255
Heather D. Stockdale, Jennifer A. Spencer, and Byron L. Blagburn
10 Foodborne Transmission.................................................................................. 289
Ynes R. Ortega and Vitaliano A. Cama
11 Waterborne: Drinking Water ........................................................................... 305
Jennifer L. Clancy and Thomas M. Hargy
12 Waterborne: Recreational Water ..................................................................... 335
Michael J. Beach
13 Waste Management........................................................................................... 371
Dwight D. Bowman
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14 Fish, Amphibians, and Reptiles ..................................................................... 387
Thaddeus K. Graczyk
15 Birds ................................................................................................................... 395
Una M. Ryan and Lihua Xiao
16 Zoo and Wild Mammals .................................................................................. 419
Olga Matos
17 Companion Animals ........................................................................................ 437
Mónica Santín and James M. Trout
18 Livestock............................................................................................................ 451
Mónica Santín and James M. Trout
19 Animal Models ................................................................................................. 485
Saul Tzipori and Giovanni Widmer
20 In Vitro Cultivation .......................................................................................... 499
Michael J. Arrowood
Index ........................................................................................................................... 527
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Preface
In the century following E.E. Tyzzer’s pioneering description of Cryptosporidium in mice, the genus
Cryptosporidium has been overlooked, rediscovered, and now found to consist of numerous species and
genotypes adapted to parasitic life in virtually all classes of vertebrates. In the past decade our knowledge
has expanded from microscopic observations of infection and environmental contamination to the
knowledge acquired from widespread application of molecular techniques to taxonomy and epidemiol-
ogy, the sequencing of the genome of two major species, and greater understanding of the biochemistry
and phylogeny of members of this fascinating genus. This second edition of Cryptosporidium and
Cryptosporidiosis has been greatly revised and expanded in response to the volume and scope of new
information on these parasites of human and veterinary importance, and the need to provide a compre-
hensive and up-to-date treatment consolidating the thousands of scientific reports.
Interest in Cryptosporidium has spread from its academic base among biologists and parasitologists
to veterinarians, physicians, epidemiologists, pharmacologists, public health specialists, drinking water
and waste water managers, swimming pool managers, farmers, backpackers, and the public in general.
Concern for prevention and treatment extends from underdeveloped communities to highly industrialized
societies, for immunocompromised persons as well as healthy populations, for persons of all ages from
infants to the elderly, and for persons caring for animals from companion animals and livestock to captive
exotic species.
Chapter 1 discusses general biological issues. It traces the history of discovery of the genus and
species, updates the taxonomy, describes the life cycle stages and their morphology, addresses host
specificity, and summarizes factors that reduce oocyst transmission. Chapter 2 introduces molecular
biology to the study of Cryptosporidium through description of data types, properties of the genome,
genetic regulation, and comparative genomics. Chapter 3 updates the biochemistry of this genus. It
describes energy and carbohydrate metabolism as well as nucleotide, fatty acid, polyamine, amino acid,
and DNA and RNA metabolism. It describes structural proteins, membrane proteins and transporters,
and delves into potential drug targets. Chapter 4 provides epidemiologists and persons interested in
transmission dynamics with detailed descriptions of outbreaks and the methods used to trace the sources.
Chapter 5 is devoted to molecular epidemiology. It describes the molecular tools, the population genetics
of Cryptosporidium species, the epidemiology of animal and human infections, and tracking sources in
water. Chapter 6 provides laboratory technicians and diagnosticians detailed descriptions of the vast
array of tests used for oocyst recovery, concentration, and purification as well as microscopic methods
for staining and observing oocysts. It summarizes immunological and molecular methods for detecting
infection and, when possible, identifying species. Chapter 7 describes host immune responses. The innate
immunity of epithelial cells and natural killer cells, T-cell-mediated immunity, parasite-specific priming
of cells, the roles of cytokines, protection from parasite specific antibodies, and vaccination against
infection are discussed. Chapter 8 provides medical workers with the description of clinical disease and
pathology in humans, including infections in immunocompetent and immunocompromised persons, and
age-related infections. It discusses organ sites, histopathology, pathophysiology, and human volunteer
studies. Chapter 9 details the vast array of compounds tested with emphasis on those found most effective
for prophylaxis and treatment of cryptosporidiosis in humans and animals. Chapter 10 reviews foodborne
transmission, the outbreaks, methods of detection in various foods, sources of contamination, decon-
tamination, and HACCP and other regulations. Chapter 11 presents the issues concerned with Cryptospo-
ridium oocysts in drinking water. These include outbreaks, methods for detection and prevention, and
government regulations related to the drinking water industry. Chapter 12 discusses factors associated
with transmission of Cryptosporidium in various types of recreational waters. It describes the outbreaks
and means to reduce or prevent future outbreaks. Chapter 13 deals with waste management, specifically:
treatment of wastewater effluent, sludge treatment, and treatment of manure from cattle and swine.
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Chapter 14 reviews infections in fish, amphibians, and reptiles, and discusses treatment, prevention, and
control. Chapter 15 addresses cryptosporidiosis in birds: the disease, immunity, prophylaxis, and treat-
ment; cultivation of the parasite; and the major species and genotypes infecting birds. Chapter 16 covers
the range of wild animals infected with Cryptosporidium from rodents, lagomorphs, insectivores, omni-
vores, ruminants, carnivores, bats, and marsupials to nonhuman primates. Chapter 17 discusses
cryptosporidiosis of major companion animal species: cats, dogs, and horses. Chapter 18 discusses
cryptosporidiosis of major livestock species: cattle, sheep, goats, pigs, and other species. Chapter 19
provides new information on animal models best suited for parasite propagation: the rodent, pig, monkey,
and gerbil models best suited for research; and models for testing parasite–host range. And finally,
Chapter 20 provides descriptions of in vitro methods of studying Cryptosporidium, the cells and media
found most useful, and the techniques for producing and storing purified parasites.
The editors acknowledge with gratitude the chapter authors for their contributions to this book.
Ronald Fayer
Lihua Xiao
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Editors
Ronald Fayer received his B.S. degree from the University of Alaska–Fairbanks and his M.S. and Ph.D.
degrees under the direction of Professor Datus Hammond at Utah State University. Dr. Fayer began his
professional career as a zoologist in the Beltsville Parasitology Laboratory of the U.S. Department of
Agriculture’s Agricultural Research Service developing in vitro cultivation methods for the protozoan
parasites Eimeria, Toxoplasma, Besnoitia, Isospora, Hepatozoon, and Sarcocystis. Following up on the
discovery of the coccidian life cycle of Sarcocystis in cell culture, studies were devoted to defining the
pathological effects in hosts including immunopathological and metabolic perturbations resulting in heart
lesions, abortions, retarded growth, and mortality. Preceding the advent of AIDS, his research emphasis
shifted to Cryptosporidium and has encompassed molecularly based surveys of prevalence in livestock;
effects of disinfectants, heat and cold on oocysts; immune responses in mouse models and calves; passive
immunotherapy with colostral antibody; drug treatments; mechanical transport hosts; identification of
new species using molecular tools; and use of molluscan shellfish as bio-indicators of fecal pollution of
surface waters. Dr. Fayer has served in a variety of administrative capacities in the Agricultural Research
Service including research leader, laboratory chief, national program leader for parasitology and toxi-
cology, director of the Animal Parasitology Institute, and assistant area director for the Northeastern
states. He has served on the editorial boards of five scientific journals and as president of the
Helminthological Society of Washington, the American Association of Veterinary Parasitologists, and
the American Society of Parasitologists. He was the recipient of a senior Fulbright Fellowship. He has
published over 325 papers in scientific journals and five books. For his contributions to research he has
received the H.B. Ward Medal from the American Society of Parasitologists, the National Distinguished
Scientist of the Year Award from the Agricultural Research Service, the Superior Service Award and the
Plow Award from the U.S. Department of Agriculture, and the Presidential Rank Award for Distinguished
Senior Professional in the career civil service.
Lihua Xiao obtained his veterinary education in China. After receiving his M.S. in veterinary parasi-
tology and teaching for 2 years at the Northeast Agricultural University in Harbin, China, he obtained
his Ph.D. in veterinary parasitology under Professor Harold Gibbs at the University of Maine and received
postdoctoral training with Professor Rupert Herd at the Ohio State University College of Veterinary
Medicine. In 1993, he moved to the Centers for Disease Control and Prevention (CDC), first as a guest
researcher, then as a senior staff fellow. He is currently a senior scientist in the Division of Parasitic
Diseases, National Center for Zoonotic, Vector-Borne, and Enteric Diseases, CDC, Atlanta, Georgia. Dr.
Xiao’s earlier research interests were mostly on the epidemiology, pathogenesis, and control of gas-
trointestinal nematodes of farm animals. Prior to the massive cryptosporidiosis outbreak in Milwaukee,
his research shifted to epidemiology and biology of cryptosporidiosis and giardiasis of farm animals.
More recently, he has focused on the taxonomy, molecular epidemiology, and environmental biology of
Cryptosporidium, Giardia, microsporidia, and other enteric protists in humans and animals, while
working simultaneously on the immunopathogenesis and vaccine development of malaria and immuno-
pathogenesis of HIV. Dr. Xiao has published over 200 scientific papers, invited reviews, and book
chapters, over half of which are on Cryptosporidium and cryptosporidiosis. He has received the James
H. Nakano citation from the National Center for Infectious Diseases, CDC, and the Outstanding Overseas
Young Scientist Award from the National Science Foundation of China. He also holds several adjunct
faculty positions at universities in the United States and China.
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Contributors
Michael J. Arrowood Centers for Disease Control and Prevention, Atlanta, Georgia
Michael J. Beach Centers for Disease Control and Prevention, Atlanta, Georgia
Byron L. Blagburn College of Veterinary Medicine, Auburn University, Auburn, Alabama
Dwight D. Bowman Department of Microbiology and Immunology, College of Veterinary Medicine,
Cornell University, Ithaca, New York
Vitaliano A. Cama Bloomberg School of Public Health, Johns Hopkins University, Baltimore,
Maryland
Jennifer L. Clancy Clancy Environmental Consultants, Inc., St. Albans, Vermont
Ronald Fayer United States Department of Agriculture, Agricultural Research Service, Animal and
Natural Resources Institute, Environmental Microbial Safety Laboratory, Beltsville, Maryland
Thaddeus K. Graczyk Bloomberg School of Public Health, Johns Hopkins University, Baltimore,
Maryland
Richard L. Guerrant Center for Global Health, Division of Infectious Diseases and International
Health, University of Virginia School of Medicine, Charlottesville, Virginia
Thomas M. Hargy Clancy Environmental Consultants, Inc., St. Albans, Vermont
Jessica C. Kissinger Center for Tropical and Emerging Global Diseases, and Department of Genetics,
University of Georgia, Athens, Georgia
Olga Matos Unit of Opportunistic Protozoa/HIV and Other Protozoa, Institute of Hygiene and Tropical
Medicine, New University of Lisbon, Portugal
Vincent McDonald Centre for Gastroenterology, Barts and the London School of Medicine, Queen
Mary College, London, United Kingdom
Gordon Nichols Centre for Infections, Health Protection Agency, London, United Kingdom
Ynes R. Ortega Center for Food Safety and Department of Food Science and Technology, University
of Georgia, Griffin, Georgia
Una M. Ryan School of Veterinary and Biomedical Sciences, Murdoch University, Perth, Australia
Mónica Santín United States Department of Agriculture, Agricultural Research Service, Animal and
Natural Resources Institute, Environmental Microbial Safety Laboratory, Beltsville, Maryland
Huw Smith Scottish Parasite Diagnostic Laboratory, Stobhill Hospital, Glasgow, Scotland
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Jennifer A. Spencer College of Veterinary Medicine, Auburn University, Auburn, Alabama
Heather D. Stockdale College of Veterinary Medicine, Auburn University, Auburn, Alabama
James M. Trout United States Department of Agriculture, Agricultural Research Service, Animal and
Natural Resources Institute, Environmental Microbial Safety Laboratory, Beltsville, Maryland
Saul Tzipori Cummings School of Veterinary Medicine, Tufts University, North Grafton,
Massachusetts
Cirle Alcantara Warren Center for Global Health, Division of Infectious Diseases and International
Health, University of Virginia School of Medicine, Charlottesville, Virginia
Giovanni Widmer Cummings School of Veterinary Medicine, Tufts University, North Grafton,
Massachusetts
Lihua Xiao Division of Parasitic Diseases, Center for Disease Control and Prevention, Atlanta, Georgia
Guan Zhu Department of Veterinary Pathobiology, College of Veterinary Medicine Biomedical
Sciences, Texas A&M University, College Station, Texas
