Fish Vaccination

Fish farming, in seawater and in freshwater, in cages, tanks or ponds, makes an ever-increasing and significant contribution to the production of aquatic food in many regions of the world. During the last few decades there has been significant progress and expansion in the aquaculture sector, characterized by intensified production and the exploitation of many new species.

Aquaculture must be a sustainable bio-production, environmentally as well as economically. Disease prevention in order to reduce losses, and the use of antimicrobials is crucial in this perspective. Vaccination has, in a few years, become the most important method for disease prevention in aquaculture, and effective prophylaxis based on stimulation of the immune system of the fish is essential for further development of the industry.

This book provides general information about disease prevention in fish by vaccination, as well as specific descriptions of the correct use of vaccines against the most important bacterial and viral infectious diseases of aquatic animals. The book is written by some of the world?s leading experts in the subject, drawn from many countries where aquaculture is a significant and expanding part of the economy.

Fish Vaccination is an encyclopedia of fish vaccinology for every present and future aquaculturist. Professionals in the aquaculture sector, including fish veterinarians and fish biologists, within the industry, in scientific institutions and regulatory authorities will all find a huge wealth of commercially important knowledge within this book. Libraries in all universities where aquaculture, biological and veterinary sciences are studied and taught should have copies of this important book on their shelves.

Contributors xv

Preface xix

1 The History of Fish Vaccination 1
Roar Gudding and Thomas Goodrich

1.1 Introduction 1

1.2 Aquaculture 2

1.3 Immunology 2

1.4 Disease Prevention 3

1.5 Scientific Production – Reviews and Conferences 4

1.6 Successes and Failures 7

1.7 The Pioneers 7

1.8 Concluding Remarks 8

References 9

2 Vaccination as a Preventive Measure 12
Roar Gudding

2.1 Introduction 12

2.2 Biosecurity and Vaccination 13

2.3 Use of Vaccination in Aquaculture 15

2.4 Vaccination Against Different Diseases 16

2.5 Herd Immunity 17

2.6 Economic Considerations 18

2.7 Risk Assessment 18

2.8 The Market for Fish Vaccines 19

References 20

3 Non-replicating Vaccines 22
Hetron Mweemba Munang’andu, Stephen Mutoloki and Øystein Evensen

3.1 Introduction 22

3.2 Classification 22

3.3 Inactivated Vaccines – Methods of Inactivation 24

3.4 Evaluation of Inactivation Efficacy 28

3.5 Measures of Efficacy for Inactivated Vaccines 28

3.6 Mechanisms of Vaccine Protection 29

3.7 Antibodies as Correlates of Protective Immunity 29

3.8 Antigen Dose as Correlate of Protective Immunity 30

References 30

4 Replicating Vaccines 33
Craig A. Shoemaker and Phillip H. Klesius

4.1 Introduction 33

4.2 Attenuation Strategies – Bacterial Vaccines 34

4.3 Attenuation Strategies – Viral Vaccines 38

4.4 Induction of Immunity 39

4.5 Vaccine Delivery 41

4.6 Vaccine Safety Considerations 41

4.7 Acknowledgement 41

References 42

5 DNA Vaccines 47
Eirik Biering and Kira Salonius

5.1 Introduction 47

5.2 Comparison of DNA Vaccines with Conventional Inactivated Products: Advantages and Disadvantages 48

5.3 DNA Vaccines for Veterinary Use 51

5.4 Biosecurity and Regulatory Considerations 52

References 54

6 Mucosal Vaccination of Fish 56
Jan H.W.M. Rombout and Viswanath Kiron

6.1 Introduction 56

6.2 History of “Mucosal” Vaccination 57

6.3 Mucosal versus Systemic Immunity in Fish 58

6.4 Immersion Vaccination 59

6.5 Oral Vaccination 60

6.6 Perspectives 64

References 64

7 Adjuvants in Fish Vaccines 68
Carolina Tafalla, Jarl Bøgwald, Roy A. Dalmo, Hetron Mweemba Munang’andu and Øystein Evensen

7.1 Introduction 68

7.2 Vaccine Formulations 69

7.3 Principles of Adjuvant Actions 70

7.4 Antigenic Component 70

7.5 Adjuvants 71

7.6 Antigen Delivery Systems 71

7.7 Delivery Vehicles 71

7.8 Emulsion Vaccines 72

7.9 Biodegradable Particulate Delivery Systems 74

7.10 Fusion Protein Delivery System 74

7.11 Immunomodulators 75

7.12 Stabilizers 81

7.13 Concluding Remarks and Perspectives 81

7.14 Acknowledgements 81

References 81

8 The Innate Immune Response in Fish 85
Jorunn B. Jørgensen

8.1 Introduction 85

8.2 Innate Immunity: A Sensing and an Effector Arm 86

8.3 Professional Phagocytes: The Macrophages and the Neutrophilic Granulocytes 86

8.4 Natural Killer (NK)-Like Cells 88

8.5 The Sensing Arm of Innate Immunity 88

8.6 TLRs are the Best Studied PRRS in Fish 89

8.7 NOD-Like and RIG-I Receptors are Found in Fish 90

8.8 Lectins are Multifunctional Sensor Molecules for Carbohydrate Ligands 91

8.9 PRRs and the Induction of Immunity 92

8.10 Cytokines in Innate Immunity 92

8.11 Interferons 94

8.12 The Complement System 95

8.13 Concluding Remarks and Perspectives 97

References 97

9 The Adaptive Immune Response in Fish 104
Stephen Mutoloki, Jorunn B. Jørgensen and Øystein Evensen

9.1 Introduction 105

9.2 Lymphocytes are the Key Cells of the Adaptive Immune System 106

9.3 Antigen Trapping and Activation of the Lymphocytes 106

9.4 Antigen Presenting Cells (APCS) of Myeloid Origin 107

9.5 Immunoglobulins and B Lymphocytes 108

9.6 T Lymphocytes 110

9.7 Cytotoxic T-Cells 111

9.8 Helper T-Cells 111

References 113

10 Development, Production and Control of Fish Vaccines 116
Dag Knappskog, Joseph Koumans, Inger Kvitvang, Arne Marius Fiskum and Rune Wiulsrød

10.1 Introduction 116

10.2 Manufacturing License 117

10.3 Vaccine Development 121

10.4 Development of Tests 122

10.5 Transfers 123

10.6 Manufacturing 124

References 127

11 Legal Requirements and Authorization of Fish Vaccines 128
Anja Holm, Byron E. Rippke and Ken Noda

11.1 Introduction 128

11.2 Manufacturer Authorization 128

11.3 Food Safety – Maximum Residue Limits 131

11.4 Genetically Modified Organisms 131

11.5 DNA Vaccines 132

11.6 Prohibition of Use of Certain Vaccines 132

11.7 Use of Vaccines that are not Authorized 132

11.8 Autogenous Vaccines 133

11.9 Regional Rules and Competent Authorities 133

11.10 The European Union and Connected EEA Countries 133

11.11 United States of America 135

11.12 Japan 137

11.13 Other Relevant Organizations: OIE, FAO, WHO 138

References 138

12 Vaccination Strategies and Procedures 140
Atle Lillehaug

12.1 Introduction 141

12.2 Timing of Vaccination 141

12.3 Water Temperature 142

12.4 Size of Fish 142

12.5 Vaccination Methods 143

12.6 Time for Protection to Develop – Duration of Protection 148

12.7 Booster Vaccination 149

12.8 Vaccination Economy 150

References 150

13 Side-Effects of Vaccination 153
Trygve T. Poppe and Erling O. Koppang

13.1 Introduction 153

13.2 Acute Side-Effects 154

13.3 Chronic Side-Effects 155

13.4 Injection Site Reactions 155

13.5 Extensive Abdominal Lesions 156

13.6 Lesions in Other Organs 158

13.7 Skeletal Lesions 159

13.8 Autoimmunity 159

13.9 Lesions in Non-Salmonid Species 159

References 160

14 Future Fish Vaccinology 162
Øystein Evensen

14.1 Molecular Technologies 162

14.2 Recombinant Vaccines 163

14.3 Marker Vaccines 166

14.4 Mucosal Vaccination 166

14.5 Vaccines Against Parasitic Diseases 167

14.6 Vaccines for Controlling Reproduction 167

14.7 Improved Formulations 168

14.8 Immunomodulation 168

14.9 Cytokines and DAMPS (Danger-Associated Molecular Pattern) as Adjuvants 169

14.10 Concluding Remarks 169

References 170

15 Vaccination against Vibriosis 172
Duncan J. Colquhoun and Atle Lillehaug

15.1 Vibriosis 172

15.2 Occurrence and Significance 173

15.3 Etiology 174

15.4 Pathogenesis 176

15.5 Vaccines 177

15.6 Vaccination Procedures 179

15.7 Vaccine Effect 179

15.8 Side-Effects 180

15.9 Regulations 180

References 181

16 Vaccination against Furunculosis 185
Paul J. Midtlyng

16.1 Introduction 185

16.2 Occurrence and Significance 186

16.3 Etiology 187

16.4 Pathogenesis and Virulence 187

16.5 Antigens 189

16.6 Vaccines 189

16.7 Vaccination Procedures 190

16.8 Effects 191

16.9 Side-Effects 192

16.10 Vaccination Against Atypical Furunculosis 192

16.11 Legal Aspects and Regulations 193

References 194

17 Vaccination against Photobacteriosis 200
Jesús L. Romalde

17.1 Occurrence and Significance 200

17.2 Etiology 202

17.3 Pathogenesis 202

17.4 Vaccines 204

17.5 Vaccination Procedures 205

17.6 Effect 206

17.7 Side-Effects 206

17.8 Regulations 207

References 207

18 Vaccination against Enteric Septicemia of Catfish 211
Phillip H. Klesius and Julia W. Pridgeon

18.1 Significance 211

18.2 Occurrence 212

18.3 Etiology 212

18.4 Pathogenesis 213

18.5 Virulence Factors 214

18.6 Vaccines and Immunity 214

18.7 Regulations (US) 220

18.8 Vaccination Practices 220

References 221

19 Vaccination against Yersiniosis 226
Andrew Bridle and Barbara Nowak

19.1 Yersiniosis 226

19.2 Occurrence and Significance 227

19.3 Etiology 227

19.4 Pathogenesis 228

19.5 Vaccines 229

19.6 Vaccination Procedures 230

19.7 Vaccine Effect 231

19.8 Side-Effects 232

19.9 Regulations 232

References 233

20 Vaccination against Streptococcosis and Lactococcosis 236
Julia W. Pridgeon and Phillip H. Klesius

20.1 Occurrence 236

20.2 Significance 236

20.3 Etiology 237

20.4 Pathogenesis 238

20.5 Vaccines 239

20.6 Vaccination Procedures and Vaccine Effect 240

20.7 Side-Effects 242

20.8 Regulations 242

References 243

21 Vaccination against Piscirickettsiosis 246
Sergio H. Marshall and Jaime A. Tobar

21.1 Occurrence and Significance 246

21.2 Etiology 247

21.3 Pathogenesis 248

21.4 Vaccines and Vaccination 248

21.5 Current Vaccine Status 249

21.6 Future Perspectives 251

References 252

22 Vaccination against Bacterial Kidney Disease 255
Diane G. Elliott, Gregory D. Wiens, K. Larry Hammell and Linda D. Rhodes

22.1 Introduction 255

22.2 Occurrence 256

22.3 Significance 256

22.4 Etiology 257

22.5 Pathogenesis 259

22.6 Vaccines 260

22.7 Vaccination Procedures 261

22.8 Vaccine Effects and Side-Effects 262

22.9 Regulations 264

22.10 Future Directions 264

References 266

23 Vaccination against Diseases Caused by Flavobacteriaceae Species 273
Krister Sundell, Eva Högfors-Rönnholm and Tom Wiklund

23.1 Introduction 273

23.2 Bacterial Gill Disease (Flavobacterium branchiophilum) 274

23.3 Columnaris Disease (Flavobacterium columnare) 275

23.4 Bacterial Cold-Water Disease (Flavobacterium psychrophilum) 278

23.5 Tenacibaculosis (Tenacibaculum maritimum) 281

References 282

24 Vaccination against Viral Hemorrhagic Septicemia and Infectious Hematopoietic Necrosis 289
Stéphane Biacchesi and Michel Brémont

24.1 Occurrence and Significance 289

24.2 Etiology 291

24.3 Pathogenesis 292

24.4 Vaccines 293

24.5 Concluding Remarks 297

24.6 Acknowledgements 297

References 298

25 Vaccination against Infectious Pancreatic Necrosis 303
Espen Rimstad

25.1 Occurrence and Significance 303

25.2 Etiology 305

25.3 Pathogenesis 306

25.4 Vaccines and Vaccine Effect 306

25.5 Vaccine-Induced Immune Responses 309

25.6 Regulations 309

References 309

26 Vaccination against Infectious Salmon Anemia 313
Knut Falk

26.1 Occurrence and Significance 313

26.2 Etiology 314

26.3 Pathogenesis 315

26.4 Vaccines 316

26.5 Regulatory Issues 317

References 318

27 Vaccination against Koi Herpesvirus Disease 321
Arnon Dishon, Ofer Ashoulin, E. Scott Weber III and Moshe Kotler

27.1 Occurrence and Significance 321

27.2 Etiology 322

27.3 Pathogenesis 323

27.4 Vaccine and Vaccination 324

27.5 Efficacy 327

27.6 Safety 329

27.7 Regulatory Issues 330

References 330

28 Vaccination against Diseases Caused by Salmonid alphavirus 334
Emilie Mérour and Michel Brémont

28.1 Occurrence and Significance 334

28.2 Etiology 335

28.3 Pathogenesis 338

28.4 Immunity and Vaccine Development 338

References 339

29 Vaccination against Diseases Caused by Betanodavirus 341
Sonal Patel and Audun H. Nerland

29.1 Viral Encephalopathy and Retinopathy (VER) 341

29.2 Occurrence and Significance 342

29.3 Etiology 342

29.4 Pathogenesis 343

29.5 Immune Status and Response to NNV 344

29.6 Vaccines 344

29.7 Replicating Vaccines 345

29.8 Inactivated Virus 347

29.9 Recombinant Protein/Peptide 347

29.10 DNA Vaccines 348

29.11 Future Prospects and Recommendations 349

References 349

30 Immunostimulation of Crustaceans 352
Indrani Karunasagar, Singaiah NaveenKumar, Biswajit Maiti and Praveen Rai

30.1 Introduction 352

30.2 Immune System of Crustaceans 353

30.3 Immunostimulants of Crustaceans 361

30.4 Acknowledgements 366

References 366

Index 373

ROAR GUDDING has been working with vaccination in veterinary medicine, including vaccination of fish, at the Norwegian Veterinary Institute for more than 25 years. He has also been Professor of Immunoprophylaxis at the Norwegian School of Veterinary Science, and is the author of textbooks about vaccination of terrestrial and aquatic animals.

ATLE LILLEHAUG has a PhD in fish vaccinology from 1993, based on field studies and clinical trials. He has been involved in the practical implementation of vaccination in Norwegian fish farming, and is now working with fish health in a broad sense. He has been the head of the Section for Fish Health at the Norwegian Veterinary Institute for several years.

ØYSTEIN EVENSEN is Professor at the Norwegian School of Veterinary Science. He has been working in the field of fish vaccination over the last 20 years. He also has more than seven years of experience from the pharmaceutical industry where he worked on the development of vaccines for fish farming globally.