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Diseases of Poultry
Diseases of Poultry
Diseases of Poultry
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Diseases of Poultry

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Diseases of Poultry is the most comprehensive reference for all aspects of poultry health and diseases, including pathogenesis, diagnostics, epidemiology, and control methods. Published in partnership with the American Association of Avian Pathologists, the Thirteenth Edition remains the international definitive reference, adding newer diagnostic methods and a new chapter on the emerging importance of zoonotic infections for poultry pathogens. Other updates include new high-quality photographs, additional discussion of conceptual operational biosecurity and disease control in organic production systems, and a greater emphasis throughout on the differences in disease incidence and treatments for the United States and other areas around the globe.

Organized logically by disease type, the book offers detailed coverage of the history, etiology, pathobiology, diagnosis, and intervention strategies, as well as the economic and public health significance, for an exhaustive list of common and uncommon diseases. Diseases of Poultry, 13th Edition is an essential purchase for poultry veterinarians, veterinary diagnosticians, poultry scientists, students specializing in poultry health, and government officials who deal with poultry health in regulatory climate.

LanguageEnglish
PublisherWiley
Release dateJul 16, 2013
ISBN9781118719732
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    Diseases of Poultry - Wiley

    Contents

    Contributors

    Preface: Historical Review of Diseases of Poultry

    Introduction

    I General Concepts of Poultry Diseases

    Chapter 1: Principles of Disease Prevention, Diagnosis, and Control

    Introduction

    Principles of Disease Prevention, Diagnosis, and Control Introduction

    Antimicrobial Therapy (Including Resistance)

    Public Health Significance of Poultry Diseases

    Chapter 2: Host Factors for Disease Resistance

    Introduction

    The Avian Immune System

    Genetics of Disease Resistance

    II Viral Diseases

    Chapter 3: Newcastle Disease, Other Avian Paramyxoviruses, and Avian Metapneumovirus Infections

    Introduction

    Newcastle Disease

    Avian Paramyxoviruses 2–11

    Avian Metapneumovirus

    Chapter 4: Infectious Bronchitis

    Chapter 5: Infectious Laryngotracheitis

    Chapter 6: Influenza

    Chapter 7: Infectious Bursal Disease

    Chapter 8: Chicken Infectious Anemia Virus and Other Circovirus Infections

    Introduction

    Chicken Infectious Anemia

    Circovirus Infections of Avian Species

    Chapter 9: Adenovirus Infections

    Introduction

    Aviadenovirus Infections

    Atadenovirus (Egg Drop Syndrome and Related Infections)

    Hemorrhagic Enteritis and Related Infections

    Quail Bronchitis

    Chapter 10: Pox

    Chapter 11: Reovirus Infections

    Introduction

    Viral Arthritis

    Other Reovirus Infections

    Chapter 12: Viral Enteric Infections

    Introduction

    Turkey Coronavirus Enteritis

    Rotavirus Infections

    Astrovirus Infections

    Avian Enterovirus-Like Virus Infections

    Enteric Parvovirus Infections of Chickens and Turkeys

    Chapter 13: Viral Infections of Waterfowl

    Introduction

    Duck Hepatitis

    Duck Virus Enteritis (Duck Plague)

    Hemorrhagic Nephritis Enteritis of Geese

    Parvovirus Infections of Waterfowl

    Chapter 14: Other Viral Infections

    Introduction

    Miscellaneous Herpesvirus Infections

    Avian Nephritis

    Arbovirus Infections

    Turkey Viral Hepatitis

    Avian Encephalomyelitis

    Avian Hepatitis E Virus Infections

    Chapter 15: Neoplastic Diseases

    Introduction

    Marek’s Disease

    Leukosis/Sarcoma Group

    Reticuloendotheliosis

    Other Tumors

    III Bacterial Diseases

    Chapter 16: Salmonella Infections

    Introduction

    Pullorum Disease and Fowl Typhoid

    Paratyphoid Infections

    Arizonosis

    Chapter 17: Campylobacteriosis

    Chapter 18: Colibacillosis

    Chapter 19: Pasteurellosis and Other Respiratory Bacterial Infections

    Introduction

    Fowl Cholera

    Riemerella anatipestifer Infection

    Ornithobacterium rhinotracheale Infection

    Bordetellosis (Turkey Coryza)

    Chapter 20: Infectious Coryza and Related Bacterial Infections

    Chapter 21: Mycoplasmosis

    Introduction

    Mycoplasma gallisepticum Infection

    Mycoplasma meleagridis Infection

    Mycoplasma synoviae Infection

    Mycoplasma iowae Infection

    Other Mycoplasmal Infections

    Chapter 22: Clostridial Diseases

    Introduction

    Ulcerative Enteritis

    Necrotic Enteritis

    Botulism

    Gangrenous Dermatitis

    Chapter 23: Other Bacterial Diseases

    Introduction

    Staphylococcosis

    Streptococcus and Enterococcus

    Erysipelas

    Avian Intestinal Spirochetosis

    Tuberculosis

    Miscellaneous and Sporadic Bacterial Infections

    Chapter 24: Avian Chlamydiosis

    IV Fungal Diseases

    Chapter 25: Fungal Infections

    Aspergillosis

    Candidiasis (Crop Mycosis)

    Sporadic Fungal Infections

    V Parasitic Diseases

    Chapter 26: External Parasites and Poultry Pests

    Chapter 27: Internal Parasites

    Introduction

    Nematodes and Acanthocephalans

    Chapter 28: Protozoal Infections

    Introduction

    Coccidiosis

    Cryptosporidiosis

    Histomoniasis (Blackhead) and Other Protozoan Diseases of the Intestinal Tract

    Miscellaneous and Sporadic Protozoal Infections

    VI Noninfectious Diseases

    Chapter 29: Nutritional Diseases

    Chapter 30: Developmental, Metabolic, and Other Noninfectious Disorders

    Chapter 31: Mycotoxicoses

    Chapter 32: Toxins and Poisons

    VII Other Diseases

    Chapter 33: Emerging Diseases and Diseases of Complex or Unknown Etiology

    Introduction

    Multicausal Respiratory Diseases

    Multicausal Enteric Diseases

    Hypoglycemia-Spiking Mortality Syndrome of Broiler Chickens

    Proventriculitis and Proventricular Dilatation of Broiler Chickens

    Index

    Advertisement

    This edition first published 2013 © 2013 by John Wiley & Sons, Inc.

    First edition, © 1943 Iowa State University Press

    Second edition, © 1948 Iowa State University Press

    Third edition, © 1952 Iowa State University Press

    Fourth edition, © 1959 Iowa State University Press

    Fifth edition, © 1965 Iowa State University Press

    Sixth edition, © 1972 Iowa State University Press

    Seventh edition, © 1978 Iowa State University Press

    Eighth edition, © 1984 Iowa State University Press

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    Tenth edition, © 1997 Iowa State University Press

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    Twelfth edition, © 2008 Blackwell Publishing Ltd.

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    Library of Congress Cataloging-in-Publication Data

    Diseases of poultry. – 13th ed. / editor-in-chief, David E. Swayne ; associate editors, John R. Glisson … [et al.].

    p. ; cm.

    Includes bibliographical references and index.

    ISBN 978-0-470-95899-5 (hardback : alk. paper)

    I. Swayne, David E.

    [DNLM: 1. Poultry Diseases. SF 995]

    SF995

    636.5′0896–dc23

    2013007104

    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.

    Cover design by Modern Alchemy LLC

    Dedicated to

    Dr. Y.M. Saif, Associate Editor,

    Diseases of Poultry, 10th edition

    Editor-in-Chief, 11th and 12th editions

    Dr. A.M. Fadly, Associate Editor,

    Diseases of Poultry, 11th and 12th editions

    Contributors

    Tahseen Abdul-Aziz

    Veterinary Pathologist

    Rollins Animal Disease Diagnostic Laboratory

    North Carolina Department of Agriculture and Consumer Services

    Raleigh, North Carolina, USA

    Claire B. Andreasen

    Associate Dean for Academic and Student Affairs

    Professor

    College of Veterinary Medicine

    Iowa State University

    Ames, Iowa, USA

    H. John Barnes

    Professor

    Department of Population Health and Pathobiology

    College of Veterinary Medicine

    North Carolina State University

    Raleigh, North Carolina, USA

    Paul A. Barrow

    Professor

    Veterinary Infectious Diseases

    School of Veterinary Medicine and Science

    Nottingham University

    Sutton Bonington, Leicestershire, United Kingdom

    Viveca Båverud

    Associate Professor, HeadDepartment of Bacteriology

    National Veterinary Institute

    Uppsala, Sweden

    Roy D. Berghaus

    Associate Professor

    Department of Population Health

    College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Alex J. Bermudez*

    Director

    Veterinary Medical Diagnostic Laboratory

    College of Veterinary Medicine

    University of Missouri

    Columbia, Missouri, USA

    Pat J. Blackall

    Senior Principal Research Fellow

    Queensland Alliance for Agriculture and Food Innovation

    The University of Queensland

    Queensland, Australia

    Gunilla Blomqvist

    Department of Virology, Immunology and Parasitology

    National Veterinary Institute

    Uppsala, Sweden

    Janet M. Bradbury

    Emeritus Professor

    School of Veterinary Science—Leahurst Campus

    University of Liverpool

    Neston, Wirral, United Kingdom

    Joseph M. Bricker

    Global Bios Research

    Zoetis

    Kalamazoo, Michigan, USA

    Bruce R. Charlton

    Professor

    California Animal Health and Food Safety Laboratory System

    University of California—Davis

    Turlock, California, USA

    Hans H. Cheng

    Supervisory Research Geneticist

    Avian Disease and Oncology Laboratory

    Agricultural Research Service

    U.S. Department of Agriculture

    East Lansing, Michigan, USA

    Richard P. Chin

    Professor

    California Animal Health and Food Safety Laboratory System

    University of California—Davis

    Tulare, California, USA

    Jens P. Christensen

    Associate Professor

    Department of Veterinary Disease Biology

    Faculty of Life Sciences

    University of Copenhagen

    Frederiksberg, Copenhagen, Denmark

    Stephen R. Collett

    Clinical Associate Professor

    Poultry Diagnostic and Research Center

    Department of Population Health

    College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Robert M. Corrigan

    Private Consultant

    Urban Rodentologist

    RMC Pest Management Consulting

    Richmond, Indiana, USA

    Rocio Crespo

    Associate Professor

    Avian Health and Food Safety Laboratory (WADDL) Department of Veterinary Microbiology and PathologyWashington State University

    Pullman, Washington, USA

    James F. Davis

    Veterinary Director of Diagnostics—North Georgia

    Georgia Poultry Laboratory Network

    Oakwood, Georgia, USA

    J. Michael Day

    Microbiologist

    Endemic Poultry Viral Diseases Research Unit

    Southeast Poultry Research Laboratory

    Agricultural Research Service

    U.S. Department of Agriculture

    Athens, Georgia, USA

    Sjaak de Wit

    Poultry Veterinarian

    Animal Health Service

    Deventer, The Netherlands

    Jean Pierre Duchatel

    Department of Infectious and Parasitic Diseases

    Faculty of Veterinary Medicine

    University of Liege

    Liege, Belgium

    Michael J. Dykstra

    Director and Research Professor

    Laboratory for Advanced Electron and Light Microscopy Methods

    Department of Population Health and Pathobiology

    College of Veterinary Medicine

    North Carolina State University

    Raleigh, North Carolina, USA

    Nicolas Eterradossi

    Agence Nationale de Securite Sanitaire de l’Alimentation, de l’Environnement et du Travail (Anses)

    Laboratoire de Ploufragan/Plouzane

    Unite de Virologie Immunologie et Parasitologie Aviaires et Cunicoles

    Laboratoire de reference de l’Organisation Mondiale de la Sante Animale pour la bursite infectieuse aviaire

    Ploufragan, France

    Aly M. Fadly

    Research Leader and Laboratory Director

    Avian Disease and Oncology Laboratory

    Agricultural Research Service

    U.S. Department of Agriculture

    East Lansing, Michigan, USA

    Naola Ferguson-Noel

    Assistant Professor

    Poultry Diagnostic and Research Center

    Department of Population Health

    College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Scott D. Fitzgerald

    Professor

    Department of Pathobiology and Diagnostic Investigation

    College of Veterinary Medicine

    Michigan State University

    East Lansing, Michigan, USA

    Steve H. Fitz-Coy

    Poultry Parasitologist

    Merck Animal Health

    Salisbury, Maryland, USA

    Richard M. Fulton

    Associate Professor

    Diagnostic Center for Population and Animal Health

    College of Veterinary Medicine

    Michigan State University

    East Lansing, Michigan, USA

    Maricarmen García

    Professor

    Poultry Diagnostic and Research Center

    Department of Population Health

    College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Richard K. Gast

    Supervisory Microbiologist

    Egg Safety and Quality Research Unit

    Agricultural Research Service

    U.S. Department of Agriculture

    Athens, Georgia, USA

    John R. Glisson

    Professor Emeritus

    Poultry Diagnostic and Research Center

    Department of Population Health

    College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Jean-Luc Guérin

    Associate Professor

    Poultry Health and Virology

    National Veterinary College of Toulouse

    Toulouse, France

    James S. Guy

    Professor

    Department of Population Health and Pathobiology

    College of Veterinary Medicine

    North Carolina State University

    Raleigh, North Carolina, USA

    Hafez M. Hafez

    Head

    Institute of Poultry Diseases

    Faculty of Veterinary Medicine

    Free University of Berlin

    Berlin, Germany

    Scott Hafner

    Veterinary Medical Officer

    Eastern Laboratory

    Food Safety Inspection Service

    U.S. Department of Agriculture

    Athens, Georgia, USA

    David J. Hampson

    Professor and Dean

    School of Veterinary and Life Sciences

    Murdoch University

    Murdoch, Western Australia, Australia

    Christopher S. Hayhow

    Director of Regulatory Affairs

    Biologics, North AmericaMerial, Inc.

    Athens, Georgia, USA

    Michael Hess

    Head

    Clinic for Avian, Reptile and Fish MedicineDepartment for Farm Animals and Veterinary Public Health

    University of Veterinary Medicine

    Vienna, Austria

    Nancy C. Hinkle

    Professor

    Department of Entomology

    College of Agricultural and Environmental Sciences

    University of Georgia

    Athens, Georgia, USA

    Fredric J. Hoerr

    Pathologist

    Veterinary Diagnostic Pathology, LLC

    Auburn, Alabama, USA

    Charles L. Hofacre

    Professor

    Poultry Diagnostic and Research Center

    Department of Population Health

    College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Tadao Imada

    One-time Research Manager (Emeritus)

    National Institute of Animal Health

    Kammondai, Tsukuba, Japan

    Sherman W. Jack

    Professor

    Department of Pathobiology and Population Medicine

    College of Veterinary Medicine

    Mississippi State University

    Mississippi State, Mississippi, USA

    Mark W. Jackwood

    Head and J. R. Glisson Professor of Veterinary Medicine

    Poultry Diagnostic and Research Center

    Department of Population Health College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Timothy J. Johnson

    Associate Professor

    Department of Veterinary and Biomedical Sciences

    College of Veterinary Medicine

    University of Minnesota

    St. Paul, Minnesota, USA

    Richard C. Jones

    Emeritus Professor

    School of Veterinary Science—Leahurst Campus

    University of Liverpool

    Neston, Wirral, United Kingdom

    Kirk C. Klasing

    Professor

    Department of Animal Science

    College of Agricultural and Environmental Sciences

    University of California—Davis

    Davis, California, USA

    Guus Koch

    Senior Scientist

    Department of Virology

    Central Veterinary Institute Wageningen UR

    Lelystad, The Netherlands

    Susan J. Lamont

    C.F. Curtiss Distinguished Professor of Agriculture and Life Sciences

    Department of Animal Science

    Iowa State University

    Ames, Iowa, USA

    Kenneth S. Latimer

    Research Clinical Pathologist

    Covance Laboratories, Inc.

    Madison, Wisconsin, USA

    David H. Ley

    Professor

    Department of Population Health and Pathobiology

    College of Veterinary Medicine

    North Carolina State University

    Raleigh, North Carolina, USA

    Catherine M. Logue

    ProfessorDepartment of Veterinary Microbiology and Preventive Medicine

    College of Veterinary Medicine

    Iowa State University

    Ames, Iowa, USA

    Larry R. McDougald

    Professor

    Department of Poultry Science

    College of Agricultural and Environmental Sciences

    University of Georgia

    Athens, Georgia, USA

    X.J. Meng

    Professor

    Department of Biomedical Sciences and Pathobiology

    College of Veterinary Medicine

    Virginia Polytechnic Institute and State University (Virginia Tech)Blacksburg, Virginia, USA

    Samia A. Metwally

    Animal Health Officer (Virologist)

    Food and Agriculture Organization of the United Nations

    Animal Health and Production Division

    Rome, Italy

    Patti J. Miller

    Veterinary Medical Officer

    Exotic and Emerging Avian Viral Diseases Research Unit

    Southeast Poultry Research Laboratory

    Agricultural Research Service

    U.S. Department of Agriculture

    Athens, Georgia, USA

    Egbert Mundt

    Animal Health

    IDT Biologika GmbH

    Am Pharmapark

    Dessau-Rossiau, Germany

    Venugopal Nair

    Head

    Avian Viral Diseases Programme

    The Pirbright Institute

    Compton Laboratory

    Compton, Berkshire, United Kingdom

    Lisa K. Nolan

    Professor and Dr. Stephen G. Juelsgaard Dean

    College of Veterinary Medicine

    Iowa State University

    Ames, Iowa, USA

    Amir H. Noormohammadi

    Associate Professor

    School of Veterinary Science

    University of Melbourne

    Werribee, Victoria, Australia

    Kenneth Opengart

    Vice President

    Live Operations and Processing

    Keystone Foods

    Huntsville, Alabama, USA

    Vilmos J. Palya

    Director

    Scientific Support and Investigation Laboratory Ceva Phylaxia Veterinary Biological Co.

    Budapest, Hungary

    Mary J. Pantin-Jackwood

    Veterinary Medical Officer

    Exotic and Emerging Avian Viral Diseases Research Unit

    Southeast Poultry Research Laboratory

    Agricultural Research Service

    U.S. Department of Agriculture

    Athens, Georgia, USA

    F. William Pierson

    Professor and Director

    Veterinary Teaching Hospital

    Virginia and Maryland Regional College of Veterinary Medicine

    Virginia Polytechnic Institute and State University (Virginia Tech)

    Blacksburg, Virginia, USA

    Silke Rautenschlein

    Professor and Director

    Clinic for Poultry

    University of Veterinary Medicine

    Hannover, Germany

    Ziv Raviv

    Assistant Professor and Head of Poultry Medicine Studies

    Koret School of Veterinary Medicine

    The Hebrew University of Jerusalem

    Rehovot, Israel

    Rodney L. Reece

    Senior Veterinary Research Officer

    NSW Department of Primary Industries

    State Veterinary Diagnostic Laboratory

    Elizabeth MacArthur Agricultural Institute

    New South Wales, Australia

    Willie M. Reed

    Dean and Professor

    College of Veterinary Medicine

    Purdue University

    West Lafayette, Indiana, USA

    Jaime A. Ruiz

    Director

    Cornell University Duck Research Laboratory

    New York State Animal Health Diagnostic Center

    College of Veterinary Medicine

    Cornell University

    Eastport, New York, USA

    Orhan Sahin

    Research Assistant Professor

    Department of Veterinary Microbiology and Preventive Medicine

    College of Veterinary Medicine

    Iowa State University

    Ames, Iowa, USA

    Y.M. Saif

    Professor and Head Emeritus

    Food Animal Health Research Program

    Ohio Agricultural Research and Development Center

    The Ohio State University

    Wooster, Ohio, USA

    Susan Sanchez

    Professor

    Athens Veterinary Diagnostic Laboratory

    College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Tirath S. Sandhu

    Director (retired)

    Cornell University Duck Research Laboratory

    College of Veterinary Medicine

    Cornell University

    Eastport, New York, USA

    Karel A. Schat

    Professor Emeritus

    Department of Microbiology and Immunology

    College of Veterinary Medicine

    Cornell University

    Ithaca, New York, USA

    Stacey L. Schultz-Cherry

    Associate Member

    Department of Infectious Diseases

    St. Jude Children’s Research Hospital

    Memphis, Tennessee, USA

    Jagdev M. Sharma

    Research Professor, Biodesign Institute

    Visiting Professor of Life Sciences, School of Life Sciences

    Arizona State University

    Tempe, Arizona, USA

    H.L. Shivaprasad

    Professor

    California Animal Health and Food Safety Laboratory System—Tulare Branch

    University of California—Davis

    Tulare, California, USA

    Leslie D. Sims

    Director

    Asia Pacific Veterinary Information Services Pty. Ltd.

    Montmorency, Victoria

    Australia

    Randall S. Singer

    Associate Professor

    Department of Veterinary and Biomedical Sciences

    College of Veterinary Medicine

    University of Minnesota

    St. Paul, Minnesota, USA

    Hanna Skarin

    Department of Bacteriology

    National Veterinary Institute

    Uppsala, Sweden

    Joan A. Smyth

    Professor

    Department of Pathobiology and Veterinary Science

    University of Connecticut

    Storrs, Connecticut, USA

    J. Glenn Songer

    Professor

    Department of Veterinary Microbiology and Preventive Medicine

    College of Veterinary Medicine

    Iowa State University

    Ames, Iowa, USA

    Edgardo Soriano-Vargas

    Professor

    Centro de Investigacion y Estudios Avanzados en Salud Animal

    Facultad de Medicina Veterinaria y Zootecnia

    Universidad Autonoma del Estado de Mexico

    Toluca, Mexico

    Stephan Spatz

    Microbiologist

    Endemic Poultry Viral Diseases Research Unit

    Southeast Poultry Research Laboratory

    Agricultural Research Service

    U.S. Department of Agriculture

    Athens, Georgia, USA

    Bruce Stewart-Brown

    Senior Vice President

    Food Safety and Quality

    Perdue Farms

    Salisbury, Maryland, USA

    David L. Suarez

    Research Leader

    Exotic and Emerging Avian Viral Diseases Research Unit

    Southeast Poultry Research Laboratory

    Agricultural Research Service

    U.S. Department of Agriculture

    Athens, Georgia, USA

    David E. Swayne

    Laboratory Director

    Southeast Poultry Research Laboratory

    Agricultural Research Service

    U.S. Department of Agriculture

    Athens, Georgia, USA

    Stephan G. Thayer

    Retired

    Poultry Diagnostic and Research Center

    Department of Population Health

    College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Daniel Todd

    Virology Branch

    Agri-Food and Biosciences Institute

    Stormont, Belfast, United Kingdom

    Deoki N. Tripathy

    Professor Emeritus

    Department of Pathobiology

    College of Veterinary Medicine

    University of Illinois

    Urbana, Illinois, USA

    Hsiang-Jung Tsai

    Professor

    School of Veterinary Medicine

    National Taiwan University

    Taipei, Taiwan

    Francisco A. Uzal

    Professor

    California Animal Health and Food Safety Laboratory System

    San Bernadino Branch

    School of Veterinary Medicine

    University of California—Davis

    San Bernadino, California, USA

    Jean-Pierre Vaillancourt

    Professor

    Department of Clinical Sciences

    Faculty of Veterinary Medicine

    University of Montreal

    St-Hyacinthe, Quebec, Canada

    Paul C.M. van Empel

    Global Technical Manager Bacteriology

    MSD-Animal Health

    Boxmeer, The Netherlands

    Daisy Vanrompay

    Professor

    Faculty of Bioscience Engineering Department of Molecular Biotechnology

    Ghent University

    Ghent, Belgium

    Vicky L. van Santen

    Professor

    Department of Pathobiology

    College of Veterinary Medicine

    Auburn University

    Auburn, Alabama, USA

    H. Vindevogel

    ProfessorClinic of Avian Diseases

    Faculty of Veterinary Medicine

    University of Liege

    Liege, Belgium

    W. Douglas Waltman

    Technical Director for Bacteriology

    Georgia Poultry Laboratory Network

    Oakwood, Georgia, USA

    Susan M. Williams

    Associate Professor

    Poultry Diagnostic and Research Center

    Department of Population Health

    College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Leslie W. Woods

    Professor

    California Animal Health and Food Safety Laboratory System

    School of Veterinary Medicine

    University of California—Davis

    Davis, California, USA

    Peter R. Woolcock

    Professor

    California Animal Health and Food Safety Laboratory System

    School of Veterinary Medicine

    University of California—Davis

    Davis, California, USA

    Guillermo Zavala

    Associate Professor

    Poultry Diagnostic and Research Center

    Department of Population Health College of Veterinary Medicine

    University of Georgia

    Athens, Georgia, USA

    Qijing Zhang

    Professor

    Department of Veterinary Microbiology and Preventive Medicine

    College of Veterinary Medicine

    Iowa State University

    Ames, Iowa, USA

    Laszlo Zsak

    Research Leader

    Endemic Poultry Viral Diseases Research Unit

    USDA ARS SAA Southeast Poultry Research Laboratory

    Agricultural Research Service

    U.S. Department of Agriculture

    Athens, Georgia

    * Deceased

    Preface: Historical Review of Diseases of Poultry

    Forewords for Diseases of Poultry, beginning with the first one written by John R. Mohler in 1943, have briefly described the nature and contents of the edition, along with substantive reasons for its publication and distribution to potential users. Mohler pointed out that for a profitable poultry industry, Knowledge of the characteristics of each disease is necessary....as the first step in building up an effective barrier against it. He further noted that This unusually comprehensive book is intended for students, veterinarians, pathologists, and workers in specialized fields. These words are as applicable today as they were 70 years ago and the importance of the text is supported by publication of 13 editions.

    For the 6th edition in 1972, Dr. P.P. Levine offered an accounting of some of the changes in the poultry industry that moved it from a small-scale farm activity to a highly sophisticated industry marketing products worth over $6 ­billion per year in the United States alone. He correctly attributed some of the many advances in disease control through eradica­tion, genetic selection, immunization prac­tices, management improvements, etc. to major advances founded in research. Such new knowledge strongly dictates a need for revised texts. Levine further predicted that infectious diseases will decline in importance; toxicologic, nutritional, genetic, and husbandry problems will demand increasing attention. Change is the order of life, and avian diseases are no exception. In the 7th edition (1978), he pointed out many of the important new advances in identifying the etiology of several conditions, and the need for Diseases of Poultry to keep up with the rapid developments in avian diseases.

    Ben Pomeroy, in the 8th (1984) and 9th (1991) editions, reiterated the need for new editions to keep up with the explosion of knowledge on the prevention and control of avian diseases. The inclusion of contributions from experts from many countries of the world, and the importance of such in the face of global issues of disease control, was emphasized by Charles Beard in the Foreword of the 10th (1997) edition. He pointed out that understanding the molecular genetics of causative agents is also important and the use of molecular methods is necessary for poultry disease researchers to understand and control infectious diseases; yet another reason for timely updates.

    The message is clear: a changing and global poultry industry and its many allied industries need the most recent information available to keep pace with the challenges of providing ­adequate health care and disease prevention. It is important not only to poultry flocks, but also to the consumers who expect safe, as well as nutritious, poultry products. This, the 13th edition, upholds the long-standing reputation of this book for keeping scientists, breeders, poultry producers, and poultry health professionals supplied with the latest and most comprehensive information available.

    Seventy years have passed since the first edition was printed. Before all details are lost forever, it is fitting to look back at how this Bible in the field of poultry diseases came to be and how it has evolved into what it is today. It all began in the 1930s. In a memorandum addressed to the American Association of Avian Pathologists (AAAP) dated December 22, 1965, H.E. Biester related the events that preceded the decision by the Iowa State College (now University) Press (ISU Press) to undertake the publication of Diseases of Poultry. During the 1930s, Louis DeVries, a member of the Department of Modern Languages at the college, translated a 1929 German book entitled Handbuch der Geflüge­lkrankheiten und der Geflügerzucht, published by Ferdinand Enke, Stuttgart. The translation laid dormant for several years until Dr. D.M. Campbell, the Chicago publisher of Veterinary Medicine, saw the translation and expressed some interest in it. Dr. Biester, who described himself as an innocent bystander, having no special interest in the project, told Dr. Campbell that the manuscript was unacceptable for a variety of reasons and he suggested that if he were serious about publication, then selected specialists should edit or rewrite the material. Dr. Biester later was pulled into the project and he ultimately concluded that the German book was obsolete. Apparently, a number of men had accepted invitations to cooperate in developing an American book, and according to Biester, they agreed that it would be better to prepare a totally new book based on American conditions.

    Thus, the die was cast. Dr. Campbell gave up his plans, and the ISU Press decided to publish an original text. Drs. Biester and DeVries served as editors, and 34 American investigators were engaged in the project. There were chapters on general subjects such as anatomy, digestion, genetics, hematology, hygiene and sanitation, nutrition, and surgery, as well as those dealing with specific infectious and noninfectious diseases and conditions. A separate chapter dealt with diseases of turkeys. In 1943, the 1st edition was ready. The publication costs were considerable for a book that was thought to have limited distribution, so it was decided to omit royalties and accept a subsidy for illustrations from the Dean of the College. Fifteen hundred copies were printed and placed on sale for $7.50; to everyone’s surprise, a second printing of 2,500 copies was needed after less than 9 months and there was yet another printing of 2,500 copies 2 years later. Royalties were then instituted! The ISU Press was concerned that without some remuneration, the authors might be reluctant to remain dedicated.

    The inclusion of Dr. DeVries as an editor is a bit puzzling since he had no medical background; perhaps it was in recognition of his effort with the translation of the German text. In any case, he was replaced in subsequent editions by Dr. L.H. Schwarte, a member of the Veterinary Research Institute in Ames who had written 4 chapters in the 1st ­edition. The book was thereafter referred to by many as Biester and Schwarte, even for a period after they were no longer associated with it. They continued at the helm through the 5th edition, published in 1965. Although Dr. Schwarte contributed several chapters in each of the first 5 editions, Dr. Biester apparently confined his efforts to editorial tasks. Their memo to the AAAP stated that they both were responsible for making the index, and they personally checked practically all of the references because they felt that they owed to the reader accuracy. A total of 61 persons served as authors under their editorial supervision; 12 of them contributed to all five editions.

    Ultimately, the passage of time dictated that Drs. Biester and Schwarte should relinquish their roles as editors and they decided that the 5th edition (1965) would be their last. As noted in the Preface to the 6th edition, it was their wish that future editions of the book become the responsibility of the AAAP… which had become a strong and representative organization to which many of the users of Diseases of Poultry belonged. Also the AAAP was already in the business of publishing the journal Avian Diseases and so it was considered a logical move. The AAAP appointed a committee, chaired by Dr. M. S. Hofstad who had been one of the book’s authors and who was on the faculty in Ames. Drs. Biester, J.E. Williams, B.S. Pomeroy, and C.F. Helmboldt filled out the committee and, in June 1966, they recommended that the AAAP sponsor future editions of Diseases of Poultry which would continue to be published by ISU Press in Ames. They asked the Board of Directors to appoint an editorial committee consisting of an editorial chairman and 4 associate editors by January 1, 1967. A letter from Dr. G.H. Snoeyenbos (AAAP secretary-treasurer) to Dr. C.A. Bottorff (AAAP president) dated November 23, 1966, suggested that Dr. P.P. Levine had declined a proposal that he assume the editorship for the book. Dr. Hofstad was subsequently named editor, and he personally requested that Drs. Helmboldt, B.W. Calnek, W.M Reid, and H.W. Yoder, Jr., be invited to be the associate editors. Each was given responsibility for a group of chapters that largely ­represented their individual interests and strengths. An agreement between the AAAP and the ISU Press was executed on May 8, 1967, and it was agreed that manuscripts would be delivered to the publisher by September 1, 1969. So the transfer was complete and official.

    The 6th edition, under totally new editorial support, underwent some significant changes. The length of the book was beginning to be of concern and there was some discussion about perhaps needing to split it into 2 volumes. To avoid this, several chapters (anatomy, nutrition, genetics, and hematology) were eliminated based on good coverage in other publications. Also, there was a consolidation of other material; for instance, all neoplastic diseases were placed in a single chapter and turkey diseases were incorporated in other chapters based on etiology. There were sweeping changes in authorship. Only 14 of the 40 contributors to the 6th edition had participated in the 5th. Clearly, a new era had arrived!

    Also, following concern for the book’s length, Dr. Hofstad asked that the number of listed references be reduced by selective citation. He agreed that the reader should find, or be directed to, all pertinent literature on each of the covered topics, the latter through citation of review papers, etc. Space allocated to references became an issue in subsequent editions as well. In the 7th and 8th editions, titles of all references were removed. This was controversial and not all editors agreed, including B.W. Calnek—senior author of this review—and reference titles appeared again beginning with his tenure as editor of the 9th edition. Interestingly, based on the number of pages, the 3rd edition (1,245 pages) was actually longer than the 11th (1,231 pages), but by increasing page size, decreasing type size, and splitting into 2 columns/page, it was possible to include more than twice the amount of written material in the latter.

    Unlike the situation with Drs. Biester and Schwarte, citations and their accuracy became the responsibility of the ­individual authors. When it was observed that many errors existed, authors of the 9th edition were asked by Dr. Calnek to check every single reference against the original work to assure accuracy. This met with an enormous number of groans and considerable resistance until each author (with perhaps an exception or 2) followed this strict instruction. The subsequent turn-around in their attitude was truly amazing when nearly all of them found errors, including the citation of references that did not even exist. It was not uncommon to detect mistakes in as many as 10% of citations in some ­chapters, probably due in large part to a common practice of copying reference citations from other lists.

    Beginning with the 9th edition, the book entered the ­electronic age. All material was submitted or copied into a word processing program that allowed spell-checking and reformatting. Initially, it was a tedious job, particularly because personal computers at that time were slow and the skill of the individual authors in mastering a new approach varied considerably. However, improvements in software and ­computers and the possibility of rapid transfer of texts between authors, editors, and the publisher made the preparation of a new edition pleasurable compared to the old hard-copy approach.

    There has been a continuum of changes that have improved Diseases of Poultry and kept it relevant over the years. For the 10th edition, the editors carefully reviewed and upgraded illustrations and for the first time included a number of color plates. Another major improvement that was gradually incorporated was the inclusion of molecular biology in many of the chapters. This was particularly important with regard to new applications of molecular techniques in diagnostic procedures, descriptions of etiological agents and significant elements of their molecular makeup, understanding the ­significance of selected genes in the pathogenesis of the ­diseases, and the development of genetically engineered vaccines. Our understanding of the fundamental nature of many diseases is now founded on the use of molecular approaches in the research laboratory.

    Another of the more significant evolutionary changes was the addition of foreign authors to make the book truly international in flavor. One of the original AAAP-appointed editors argued strongly that Diseases of Poultry should be an American book, and the authorship was so aligned. The 6th edition had the first foreign contributor, although she (Bela Tumova, from Prague, Czechoslovakia) was actually a visiting professor at the University of Wisconsin working with B.C. Easterday on avian influenza at the time. It wasn’t until the 8th edition that invitations to contribute to the book were extended to workers outside of the United States. Drs. P.M. Biggs and L.N. Payne from England and Drs. J.B. McFerran and M.S. McNulty from Northern Ireland thus paved the way by providing parts of the chapters on neoplastic diseases, adenoviruses and miscellaneous viral infections. The next edition (9th) was truly international with 17 contributors from 8 countries outside of the U.S., and by the 11th edition, there had been a total of 25 different contributors from 11 countries. The worldwide reputation the book was certainly enhanced by the selection of authors based on their knowledge and contributions to our understanding of individual diseases and conditions without regard to their geographic location.

    The importance of Diseases of Poultry as a text for the world is also reflected in its translation into foreign languages or publication in a copied form in other countries. There have been several authorized translations into Spanish, Chinese, and Russian, and an agreement between the publisher and India has allowed what is essentially a photocopied version of the original to be made.

    An ongoing review of the relative importance of individual diseases or conditions has led to a good deal of reshuffling over the years. Chapters have been added, combined, split, or eliminated to meet the changing picture of what is important to the field of avian diseases and disorders. Periodically, and especially with a change in authorship, major rewriting of some sections takes place. New chapters such as one dealing with new and emerging diseases appear when needed.

    Beginning with the 12th edition emphasis was placed on the significance of each disease to public health considering the ever increasing interest in food safety. Emphasis was also placed on standardizing the format for all the subchapters by using the same headings thus making the book more reader friendly. In addition several non-USA residents became contributing authors.

    Despite our increased understanding of disease processes there remains much to learn. Only in very few cases do we know the molecular markers of virulence, pathogenicity, or immunogenicity of pathogens and, needless to say, the molecular basis for disease resistance is a wide open field. In many cases, our understanding of disease-triggering mechanisms is poor and diseases of multiple etiologies remain problematic.

    As indicated, the introduction of molecular techniques has greatly enhanced our understanding of disease but it has also created some confusion as we strive to understand the underlying genetics of important biologic characteristics of pathogens. The early euphoria from thinking that 1 gene is responsible for a given biologic characteristic is being replaced by the realization that more than 1 gene is usually involved. This makes it difficult to decipher our observations. Another point of confusion resulted from the definition of the term genotype and early attempts to relate it to serotype or protective type. Again, it became clear that such a relationship is lacking in most cases and genotyping is useful mainly for epidemiologic studies. Considering all these gaps in our knowledge, and the unprecedented speed of knowledge generation, it is understandable that we need a new edition every 5–6 years to keep our text continually updated.

    Finally, the euphoria of the last century, suggesting that we will conquer infectious diseases as stated to the U.S. Congress in 1969, by then Surgeon General of the United States William H. Stewart (We can close the books on infectious diseases…), and our own P.P. Levine’s prediction in 1973 stated earlier in this section (Infectious diseases will decline in importance…) proved terribly wrong. Microbes are a tough and nimble foe capable of changing and adjusting to new environments mostly created by man. Thus, we think that infectious diseases will continue to be a top health priority for poultry, another reason why we continue to need new editions of this book.

    Changes in editors occurred over the years so that by the 11th edition, none of the 1968 group appointed by the AAAP remained. After riding herd on 3 editions (6th–8th), Dr. Hofstad retired and was replaced by Dr. Calnek (9th and 10th) and he, in turn, passed the baton to Dr. Y.M. Saif ­(11th–12th), and most recently to Dr. D.E. Swayne (13th). Likewise, associate editors that have replaced or been added to the original group appointed by the AAAP in 1968 include Drs. H.J. Barnes (8th–11th), C.W. Beard (9th and 10th), L.R. McDougald (10th–13th), Y.M. Saif (10th), J.R. Glisson (11th–13th), A.M. Fadly (11th and 12th), D.E. Swayne (11th and 12th), L.K. Nolan (12th and 13th), and D.L. Suarez and V. Nair (13th). Dr Nair is the first non-USA resident to serve on the editorial board of the book, another indication of the universal reach of the book.

    In summary, it is obvious that the Bible in the field of avian diseases is an evolving, vibrant, and ever-current source of information relevant to all practitioners in the field of poultry medicine. It continues to be a reference source of ­significance to a vast number of persons with many different relationships to the poultry industry.

    Bruce W. Calnek

    Y.M. Saif

    David E. Swayne

    Introduction

    This edition is appropriately dedicated to 2 colleagues who have devoted their careers to creating new knowledge and passing on existing wisdom on poultry health through successive editions of Diseases of Poultry: Dr. Y.M. Saif, who served as Associate Editor for the 10th edition and Editor-in-Chief for the 11th and 12th editions, and Dr. Aly Fadly, who served as an Associate Editor for the 11th and 12th editions. Both have been instrumental in maintaining the high quality of this book, and, indeed, we are highly grateful for their efforts. In addition, I am personally grateful to Dr. Saif, who has mentored me over the past quarter century as a scientist, writer, and editor, especially in the past 5 years in transitioning to Editor-in-Chief for the 13th edition of Diseases of Poultry. With this edition, Drs. Venugopal Nair and David L. Suarez joined the existing Associate Editors, Drs. John R. Glisson, Larry R. McDougald, and Lisa K. Nolan. We are very appreciative of their services.

    This edition has a major emphasis on electronic versions with expansion of color figures and availability of 3 specific electronic formats to accommodate a variety of media from traditional computers to stand-alone electronic books to smartphones. The new generation of poultry veterinarians and scientist are living and working in the electronic age and rely upon instantaneous access to crucial information to do their jobs, and Diseases of Poultry will be at their fingertips or in their pockets for daily and mobile use. In addition, the widely used high quality, hard copy is also preserved as a reference text, but with fewer color plates than in the electronic formats.

    This 13th edition represents a continuation of the tradition established earlier of providing the latest information on poultry diseases. Earlier trends of expansion of authorship to include authors from around the globe were continued in this edition, as was the appointment of our first non-USA Associate Editor. The Preface was updated and expanded to a more comprehensive review of the history of Diseases of Poultry, including valuable new metric data.

    All of the book chapters were updated with the most current and accurate knowledge and many with new figures, especially color figures of gross lesions. Much of the historical information was removed and readers are referred to prior editions. A new subchapter on public health significance of poultry diseases was added to Principles of Disease Prevention, Diagnosis, and Control, (Chapter 1) and public health information was reduced in the individual chapters. A new subchapter on enteric parvoviruses was added to Chapter 12, Viral Enteric Infections. Major revisions were accomplished for Principles of Poultry Production, Health and Disease Control in Commercial Systems; Newcastle Disease; Avian Paramyxoviruses 2-11, and Internal Parasites. With no new cases in 2 decades, the torovirus chapter was removed and readers are referred to prior editions of Diseases of Poultry for such information.

    Subchapters on Dermal Squamous Cell Carcinoma and Multicentric Histiocytosis were merged into the subchapter on Other Tumors. Cochlo­soma anatis Infections was merged in with the subchapter on Miscellaneous and Sporadic Protozoal Infections. These changes were dictated by the increasing or decreasing significance of some diseases or the increasing knowledge on a given disease. Some subchapters have been moved to different chapters because of recent findings indicating that they fit within different areas.

    A sincere thank you goes to the authors who contributed to the current and earlier editions of the book. It has been a wonderful experience working with all of you.

    The personnel at Wiley-Blackwell that worked on this ­edition have been most helpful and accommodating, and we sincerely appreciate their support.

    This is the first time that I served as Editor-in-Chief of Diseases of Poultry, and I (DES) am indebted to my colleagues, the associate editors, for their tireless efforts in the review process and their support and advice.

    Finally, I would like to acknowledge special assistance provided by two students, Jessica Self and Jamie Conner, in reviewing and formatting reference lists and retyping figure legends.

    Editor-in-Chief

    David E. Swayne

    Associate Editors

    John R. Glisson

    Larry R. McDougald

    Venugopal Nair

    Lisa K. Nolan

    David L. Suarez

    I  General Concepts of Poultry Diseases

    Chapter 1  Principles of Disease Prevention, Diagnosis, and Control

    Chapter 2  Host Factors for Disease Resistance

    Chapter 1

    Principles of Disease Prevention, Diagnosis, and Control

    Introduction

    Stephen R. Collett

    Poultry product prices have not kept pace with rapidly escalating input costs, causing profit margins to diminish throughout the poultry industries. Producers are faced with the challenge of consistently achieving expected performance while satisfying stringent food safety and animal welfare requirements; they can no longer tolerate even the slightest deviation from expected performance.

    In an effort to cost-effectively meet demand, producers have had to increase the size and throughput of their production systems. These large close confinement rearing systems, designed to improve economies of scale and maximize productivity by optimizing bird comfort, unfortunately also increase the risk of disease challenge. The physiological stress of keeping pace with genetic potential for production makes birds more vulnerable to disease challenge, and the close proximity of susceptible hosts increases the chance and rate of infectious disease spread. Diseases previously recognized as unimportant, because they have been adequately controlled, have now reemerged as significant concerns. Most of today’s disease challenges are not new problems; rather, they have merely expanded their geographic distribution or reemerged primarily because of management techniques and production system design constraints.

    Methods of disease control have evolved with intensification of the industry. While initially focused on diseases of catastrophic nature, attention has rapidly shifted from defined, clinical disease at the individual house or farm level to less well-defined subclinical disease and bird welfare. Similarly, cost justification in decisions on whether or not to implement prevention or control measures has become more complex, requiring the aid of formal economic appraisal.

    To maximize flock efficiency, disease challenge management requires a carefully designed, multi-tiered approach, which includes consideration of elements ranging from flock health and productivity to chick or poult viability. In addition, the term efficiency implies the all-important economic component. Because the production system is profit driven, decisions regarding disease challenge can rarely be made based solely on biological grounds. Disease management intervention also requires sound economic justification, which begins with clearly defining the financial risk associated with disease.

    Unless a disease poses a specific risk to human health or animal welfare, its mere presence in a flock may not be significant from a business perspective. It is often difficult for the veterinarian, trained in disease prevention, diagnosis, and control, to appreciate that the presence of a disease in a flock could be considered superfluous. Unless it is economically advantageous to take action against a disease challenge, its presence in a flock is tolerated. Intervention strategies are consequently chosen based on both their economic and biological efficiency. This process requires a dynamic, integrated combination of an epidemiologic and economic analysis to determine and quantify the production effect of the disease challenge as well as a proposed intervention strategy. Such integrated analysis has become far more significant in today’s intensive production systems because the outcome of disease challenge is so markedly influenced by the environmental conditions.

    The economic impact of disease is difficult to assess. This is particularly so in a production system in which the economic return is governed not only by flock productivity but also by product quality and viability. In addition, the consequential loss from disease challenge will be, at best, only partially recoverable. Using the cost of disease to justify intervention over-emphasizes the consequence of inaction and it is only useful in justifying intervention strategies directed at disease prevention. As the process of economic analysis has evolved, the focus has shifted from the cost of disease to the benefit derived from disease control strategies.

    Bird performance continues to increase linearly because of efficient breeding and selection programs, but each increment in improvement is 1 step closer to the point of physiological limit and diminishing returns. Production systems need to be modified to better satisfy bird requirements, and disease prevention, diagnosis, and control strategies changed to preclude physiological, nutritional, and agent-induced pathologies from affecting performance.

    Acknowledgement

    The author is greatly indebted to Alex J. Bermudez for his contribution to earlier editions of this subchapter.

    Principles of Disease Prevention, Diagnosis, and Control Introduction

    Stephen R. Collett

    Flock Health

    Disease is the antithesis of health but neither state is easy to define in production animals. Health is defined in the human individual as a state of physical, mental, and spiritual well-being. It is impossible to apply this definition to an animal, and production animals have in the past been classified as healthy if they were free from clinical disease and performing to standard. Although individual animals are frequently described as healthy or diseased, these terms are not mutually exclusive. The impact of disease challenge on productivity is apparent long before clinical signs of disease appear. Production animals are expected to perform at their genetic potential, and to achieve this they need to be physically and mentally well or stress free.

    Stress has been defined as a nonspecific response of the body to any demand made upon it. From a physiological point of view this can be restated as the metabolic response of the body to external factors that impact well-being (34). Stress is cumulative and only measurably impacts performance once the aggregate of each individual stress exceeds the host’s ­coping mechanisms. An interesting study by Klasing et al. (52) has shown that the degree to which an adverse stimulus or stress will negatively impact bird performance is directly proportional to the existing stress load. Any stress will impact productivity once the stress threshold is surpassed. In a production system in which animals are expected to produce at genetic potential, the definition of health needs to be expanded to freedom from disease or stress. In this context, health is proportional to the difference between stress level and stress threshold.

    Disease prevention and control strategies tend to be too focused on addressing the precipitating cause, and too little attention is given to the predisposing causes of disease. The traditional paradigm of disease has been shaped by the study of specific diseases in individual animals and tends to overemphasize the importance of the infectious agent. In intensive animal agriculture it is the influence of environmental disease determinants that decides the economic outcome of infectious agent challenge. The focus of flock health management has consequently shifted. Initially aimed at avoiding mortality because of an inadequate immune response, health management is now directed at avoiding an exaggerated or inappropriate immune response because it may depress productivity. The task of the veterinarian has shifted from the prevention, diagnosis, and control of specific disease conditions in the individual bird to preventing and limiting the consequence of more complex multifactorial disease outcomes to maximize the productivity of the flock.

    Resistance and Resilience

    An animal’s resistance to disease can be defined as its capacity to prevent an overwhelming infection by a disease-causing organism. Disease resistance is determined by immune competence and health status at the time of challenge. Because stress negatively impacts health it also negatively impacts resistance. Ironically, the process of mounting an effective immune response is itself a source of stress because of the demands made on the immune system and the consequence of the resulting fever response. An immune response that is adequate to contain disease can be considered as the cost of health. There is a delicate balance between too little and too much because an inappropriate immune response, whether inadequate or excessive, will depress performance.

    The resilience of an animal is a measure of its capacity to continue to perform while preventing a disease challenge from causing an overwhelming infection. As with resistance, resilience is negatively impacted by poor health but in this case the negative impact of the resulting stress is more significant. The chemical messengers (cytokines) released in response to a disease challenge depress production directly by influencing metabolism and indirectly by suppressing appetite and feed intake (52). While immune response is crucial to maintaining health, its consequence is depressed productivity.

    The skin and respiratory, urogenital, and gastrointestinal tracts form the interface between foreign (antigenic) material and animal cells (self). To protect the bird from disease the immune system has to develop exquisite sensitivity as to whether foreign antigens are friend (nutrients or normal flora) or foe (pathogenic). An inappropriate immune response to gastrointestinal antigens will, for example, have a negative impact on feed efficiency. The fever response induced by foreign antigens will depress feed intake, while the inflammatory response damages the gut lining, thus reducing the nutrients available for production. The capacity of an animal to fight off a disease challenge while avoiding the negative impact of the induced immune response on productivity (resilience) depends on how close the prevailing level of stress is to the bird’s stress threshold. The success of any health program thus hinges on balancing immunity and health to maximize resilience. There is a dynamic interface between nutrition, immunity, and productivity. The aim of any production veterinarian should be to optimize feed utilization by modulating the immune response: enhancing the protective response to prevent clinical disease while simultaneously suppressing the acute phase or fever response.

    Population Dynamics

    Like human medicine, traditional veterinary medicine is focused on the study of the disease process in individuals. In modern flock medicine, where the emphasis is on prevention, diagnosis, and control of disease in finite and confined populations, the focus shifts to the epidemiology of the disease. Flock health is a rather nebulous term. Because health and disease are not mutually exclusive, individual birds within the flock will at any point in time be in various stages of health/disease (Poisson distribution). At what point is a flock diseased or healthy? Productivity gives a good estimate of an individual’s state of well-being and welfare. Similarly, a flock that is performing to standard is assumed to be healthy, based on the fact that the birds act and produce as an equivalent nonstressed sibling would in a laboratory situation. This approach unfortunately takes little cognizance of the flock variance, because flock performance indicators are based on flock averages. Population variance or range is a much better indicator of flock health.

    Modern population-based health management requires the complex integration of animal husbandry (housing environment), medical management (host and agent factors), and epidemiological practice (analysis of causal relationships). Modern risk management approaches facilitate the transformation of epidemiologically derived statistical data into a population health management tool. The probability of exposure to health risk factors and their expected outcomes are used first, to guide the response to a disease challenge, and second to improve the health program design/management to better respond to future potential population health challenges.

    Each component of the production process can be evaluated on the basis of cost and contribution. In the past, intensive agriculture has been production driven, and contribution measured in terms of performance. In today’s market-driven enterprise, value is regarded as a function of quality, yield, and cost of production with the emphasis shifting from performance to profit through the chain of realization. In this scenario the simplest strategy for improving productivity is to reduce within- and between-flock variance. By reducing variability and thus eliminating the extremes, it is possible to improve the quality, speed, and cost of production. Improved uniformity translates to improved productivity and hence profitability. Health (the difference between stress level and stress threshold) is probably the single most important determinant of flock uniformity. Within a group of animals the threshold and level of stress experienced by each individual will vary. The relative efficiency of a production manager to minimize in-house environmental variation, and therefore host-, agent-, and environment-dependent stress, is reflected in flock uniformity.

    Challenges of Disease Prevention, Diagnosis, and Control in Modern Poultry Production

    Because the goal of a poultry operation is to convert feed into food as economically as possible, it is critical to manage both the risk and consequence of disease challenge. While the biological efficiency of feed conversion is governed primarily by intrinsic or genetic determinants, in an intensive production system it is the extrinsic disease determinants that ultimately decide the efficiency of the operation in both biological and financial terms. Capital investment in the housing’s environmental control capability, and the effective operation of these controls, is fundamental to economic success. Even subtle disease challenge such as vaccination with live respiratory disease vaccines can compromise efficiency if exacerbated by environmental disease determinants.

    Viral diseases are challenging to control because there are no effective treatment options, while bacterial, protozoal, and parasitic diseases present a challenge because the treatment options are either no longer available or no longer effective. The approach to controlling diseases within these 2 categories is very different.

    The molecular structure of a virus particle is relatively simple, making immunological recognition very acute and the control of known viral diseases possible through immunization. Provided the immune system has been primed by vaccination, immunological protection against viral disease challenge is usually highly successful. Emerging and reemerging viral diseases arise when novel or immunologically distinct viruses are introduced into naïve populations (48). In the absence of prior exposure, immune recognition and activation is delayed and the extent of the primary immune response is frequently inadequate to prevent clinical disease (45). Under such conditions virus replication and spread occurs rapidly with potentially devastating consequences (22). While the majority of emerging viral diseases in humans are the result of exposure to novel viruses, it is the emergence of variant strains that pose the biggest threat to the poultry industry (88). Although controlled environment housing and good biosecurity practices have been highly effective in preventing the introduction of novel viruses, increased population density and vaccination have likely enhanced the emergence of variant strain viruses. The high population densities provide the opportunity for antigenic shift through gene recombination, while vaccination creates positive selection pressure for the variant strain viruses (79).

    In contrast to viruses, bacteria and protozoa are structurally and immunologically complex, making protection through vaccination much less successful. Although a great deal of research effort is and has been focused on developing effective immunization strategies for these diseases, antibiotics have remained the primary means of control (24)—a point well illustrated by the escalating difficulties experienced in the European Union with the current systematic withdrawal of in-feed antibiotics (33). It is no coincidence that the downward trend in prophylactic (in-feed) antibiotic usage has been matched by an increase in therapeutic use (58). Many expert committees blame the use of in-feed antibiotics in food animal agriculture for the proliferation of antibiotic-resistant strains of bacteria and for the increase in prevalence of antibiotic-resistant infections in humans (47). This is undoubtedly providing the impetus to ban in-feed antibiotic use, even though a link to increased antibiotic-resistant bacterial disease in humans has not been conclusively established (29). Consumer pressure to remove antibiotics from the food animal nutritionist’s arsenal is, however, winning the battle and the trend toward reemergence of previously controlled bacterial and protozoal diseases will likely continue. The industry must adapt to remain competitive.

    The Principles

    Disease prevention and control involves the 3 interrelated processes of bioexclusion, surveillance, and biocontainment. Disease prevention is difficult, expensive, and requires total commitment because it invariably involves eradication. Eradication programs are appropriate when the economic consequence of the disease is so devastating that it is economically advantageous to implement such drastic control measures. It is only feasible if there is an effective means of detecting infection, containing the infection through clean-out and disinfection, and preventing dissemination of the disease causing agent (75). There are 3 categories of disease for which eradication is an appropriate means of control: those that significantly threaten public health, those that have a devastating effect on bird performance, and those that severely compromise product quality. With diseases of this nature, the control effort is focused on the complete elimination of the agent from the environment (75). This places the emphasis on preventing contact between the agent and the host (bioexclusion). Early diagnosis and containment is in this case the contingency plan for failure in bioexclusion.

    In contrast to eradication, control programs are aimed at limiting disease challenge to a tolerable level. There is a subtle shift in emphasis from prevention, through bioexclusion, early detection, and elimination, to reducing the consequence or ­economic impact of the disease, i.e. damage control. Although monitoring and surveillance are still used to gather prevalence data, the primary focus is to measure the level of protection and challenge, not the mere presence of the disease. The principles of prevention through biosecurity still apply, but in a disease control program the focus shifts to limiting the extent and consequence of exposure. In reality, many of the biosecurity measures taken to eradicate the more devastating diseases provide a solid foundation for the control of the erosive diseases, and immunization is usually used to bolster host resistance.

    Methods of disease management have evolved with intensification of the industry. While initially focused on diseases of catastrophic nature, focus has rapidly shifted from defined, clinical disease at the individual house or farm level to less well-defined sub-clinical disease and bird well-being or welfare. Similarly, cost justification in decisions on whether or not to implement control measures has become more complex, requiring the aid of formal economic appraisal.

    The presence of disease in a flock only becomes significant when the functional derangement of normal metabolic and homeostatic processes causes a decline in productivity that is sufficiently large to affect economic efficiency. This occurs either as a result of disease-induced anorexia or through specific effects on the physiological processes of nutrient metabolism, respiration, and excretion. Only in severe cases does disease cause mortality, and yet mortality rate is used universally as a measure of flock health status. Health and disease are not mutually exclusive, yet a flock is assumed to be healthy when it is performing to standard and is free of clinical disease. Productivity provides a much more sensitive measure of flock health, but because it is the composite output of a population it gives no indication of the variance in health status between individuals within the flock. Within-flock variance provides the most sensitive measure of flock health status but flock uniformity is still a fairly coarse measure of health status.

    Disease challenge management must be considered to be an integral part of any poultry business risk management program. It involves the development and implementation of a stringent biosecurity plan which comprises a hierarchy of components directed at preventing or limiting the risk and consequence of disease. Economic analysis is a critical step in biosecurity plan design because resource allocation must match risk (10). Although it is difficult to accurately determine the precise risk and consequence of a disease challenge, it is possible to rank disease challenge according to relative risk (39). From a risk analysis point

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