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FINFISH AQUACULTURE DIVERSIFICATION

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Page 1: FINFISH AQUACULTURE DIVERSIFICATION20Finfish... · 12.4.1 Atlantic salmon, Salmo salar 255 12.4.1.1 Life cycle of Atlantic salmon 257 12.4.1.2 Farming of Atlantic salmon 261 12.4.1.3

FINFISH AQUACULTURE DIVERSIFICATION

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We would like to honour the memory of our eminent colleague, Dr Joseph A. Brown, whose enthusiasm for aquaculture diversification

provided inspiration for the book. He was greatly appreciated for his kindness, accessibility, involvement and impact on a wide range of endeavours.

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FINFISH AQUACULTURE

DIVERSIFICATION

EDITED BY

Nathalie R. Le FrançoisBiodôme de Montréal, Québec, Canada

Malcolm JoblingUniversity of Tromsø, Norway

Chris CarterUniversity of Tasmania, Australia

Pierre U. BlierUniversité du Québec à Rimouski, Québec, Canada

ASSISTANT EDITOR AND ILLUSTRATOR

Arianne SavoieUniversité du Québec à Rimouski, Québec, Canada

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CABI is a trading name of CAB International

CABI Head Office CABI North American OfficeNosworthy Way 875 Massachusetts AvenueWallingford 7th FloorOxfordshire OX10 8DE Cambridge, MA 02139UK USA

Tel: + 44 (0)1491 832111 Tel: + 1 617 395 4056Fax: + 44 (0)1491 833508 Fax: + 1 617 354 6875E-mail: [email protected] E-mail: [email protected]: www.cabi.org

© CAB International 2010. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopy-ing, recording or otherwise, without the prior permission of the copyright owners.

A catalogue record for this book is available from the British Library, London, UK.

Library of Congress Cataloging-in-Publication Data

Finfish aquaculture diversification / edited by Nathalie R. Le François … [et al.]. p. cm. Includes bibliographical references and index. ISBN 978-1-84593-494-1 (alk. paper)1. Fish culture. I. Le François, Nathalie R. II. Title.

SH151.F46 2010639.3--dc22 2009015984ISBN-13: 978 1 84593 494 1

Commissioning editor: Rachel Cutts Production editor: Fiona Harrison

Typeset by SPi, Pondicherry, India.Printed and bound in the UK by the MPG Books Group.

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v

Contributors xviiAcknowledgements xxi

I Aquaculture Diversifi cation: an Introduction 1

1 Fish Culture: Achievements and Challenges 3Malcolm Jobling

1.1 Introduction 31.2 Application of Emerging Technologies to 4

Aquaculture Research1.3 Stock Improvement 6

1.3.1 Sex-reversal and production of monosex 7populations and triploids

1.3.2 Induction of sterility 81.3.3 Interactions between farmed and wild fish 91.3.4 Transgenic fish 10

1.4 Feed Ingredients and Resources 111.4.1 Feed ingredients of plant origin 12

1.4.1.1 Transgenic plants 121.5 Health Management and Disease Control 15

1.5.1 Diagnostics and disease treatment 161.5.1.1 Vaccines 171.5.1.2 Antibiotics and other 17

chemotherapeutic agents1.6 Food Products: Nutritional Composition, Food Safety 18

and Traceability1.6.1 Analysis of aquaculture products 19

Contents

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vi Contents

1.7 Fish as Food: Consumer Attitudes (and the Question 21of GM-Fish)

1.8 Concluding Remarks 23

2 Fish Culture: the Rearing Environment 33Malcolm Jobling

2.1 Introduction 332.2 Water Sources and Their Consequences for 34

Rearing Systems2.2.1 Ponds 342.2.2 Cages and pens 362.2.3 Land-based systems 36

2.3 Biosecurity: a Key for Disease Prevention 392.4 Water Quality 40

2.4.1 Salinity 412.4.2 Oxygen 442.4.3 Nitrogenous products: ammonia, nitrite and nitrate 45

2.5 Temperature 492.5.1 Temperature, reproduction and development 51

2.6 Light 532.7 Fish Welfare Under Culture Conditions 55

3 Fish Culture: Feeds and Feeding 61Malcolm Jobling

3.1 Introduction 613.2 Nutrients and Nutritional Requirements 62

3.2.1 Metabolism and body maintenance: fish versus 64terrestrial livestock

3.3 Aqua-feeds for Farming Fish 663.3.1 Feeds for different life history stages 673.3.2 Fishmeal and fish oil: a reduced reliance on these 70

feed ingredients3.3.2.1 Organic contaminants in fish and 71

fish feeds3.3.2.2 Physiological effects of organic 72

contaminants3.3.2.3 Fishmeals as a protein source in 73

aqua-feeds3.3.2.4 ‘Alternative’ protein sources in 74

aqua-feeds3.3.2.5 The need for marine fish oils in 75

aqua-feeds3.4 Feeding Behaviour, Feeding Routines and Feed 77

Delivery Systems3.4.1 Feeding methods and routines: assessment 77

of satiation

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Contents vii

3.4.2 Automatic feeding systems 793.5 Environmental Protection and Related Issues 82

4 Farmed Species and Their Characteristics 88Malcolm Jobling

4.1 Introduction 884.2 Characteristics Desirable in a Farmed Species 894.3 Fish as Food: A Comparison with Terrestrial Livestock 914.4 Natural Diets of Farmed Animals 924.5 Domestication and Genetic Selection 934.6 Culture Species 944.7 Concluding Comments 96

5 Considerations for the Selection and Commercialization 100of New or Alternate SpeciesNathalie R. Le François, David Otton and Guillaume Werstink

5.1 Introduction 1005.2 Why Develop a New or an Alternate Species? 1015.3 Who Is Likely To Develop New Species and Products? 1035.4 When Is This Development Likely To Occur? 1055.5 What Does the Development Process Entail? 1055.6 How Should Commercialization Be Conducted? 1065.7 Conclusion 112

6 A Systematic Market Approach to Species Diversifi cation: 115a French Case StudyMarc Suquet

6.1 Introduction 1156.2 Methodology 118

6.2.1 Enquiries 1186.2.2 Selection procedure 118

6.3 Constraints and Limitations of the Model 1196.4 Case Study: The Case of the French Atlantic, the 123

Channel and the North Sea Coasts6.4.1 Enquiries 1236.4.2 Selection procedure 125

7 The Agribusiness Approach 132David Otton and Lawrence Dooley

7.1 The Concept of Agribusiness 1327.2 Method 1327.3 The Agribusiness System 1327.4 Case Studies: Catfish, Atlantic Salmon and Barramundi 136

7.4.1 Channel catfish (Ictalurus punctatus) 1377.4.2 Atlantic salmon (Salmo salar) 138

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viii Contents

7.4.3 Barramundi (Lates calcarifer) 1397.5 Conclusion 1407.6 Future Trends and Issues 140

II Finfi sh Species Description 145and Biotechnical Analysis

8 Quantitative Approaches for Identifying Finfi sh Species 147Suited for Sustainable and Productive AquacultureNathalie R. Le François, Arianne Savoie and Pierre U. Blier

8.1 Introduction 1478.2 Site-specific Selection of the Most Suitable Species and 149

Production Strategy: a Québec (Canada) Case Study8.2.1 Collection of species-specific biotechnical data 1508.2.2 Description of the model 150

8.2.2.1 Step 1 1508.2.2.2 Step 2 1538.2.2.3 Step 3 153

8.2.3 Results and discussion 1538.2.4 Particularities, constraints and limitations 154

of the model8.3 Evaluation of Commercial Species’ Attributes in Terms of 155

Productivity/Sustainability8.3.1 Collection of species-specific biotechnical data 1568.3.2 Criteria selection and description 157

8.3.2.1 Reproduction and incubation 1578.3.2.2 Larval and juvenile production 1608.3.2.3 On-growing and commercialization 161

8.3.3 Results and discussion of cold, temperate and 165warmwater fish species analysis

8.3.3.1 Coldwater fish species 1658.3.3.2 Temperate fish species 1698.3.3.3 Warmwater fish species 172

8.3.4 Particularities, constraints and limitations of 173the model

9 The Sturgeons (Family: Acipenseridae) 178 Matthew Litvak

9.1 General Introduction 1789.2 Farming of Sturgeons 1799.3 Shortnose Sturgeon, Acipenser brevirostrum 180

9.3.1 Biology and culture 1809.3.1.1 Gross anatomy 1819.3.1.2 Distribution 1829.3.1.3 General life history 1829.3.1.4 Behaviour/adaptability 185

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Contents ix

9.3.1.5 Fishery status 1859.3.2 Broodstock management and hatchery operations 1859.3.3 On-growing to market size 187

9.4 Atlantic Sturgeon, Acipencer oxyrinchus oxyrinchus 1889.4.1 Biology and culture 188

9.4.1.1 Gross anatomy 1889.4.1.2 Distribution 1899.4.1.3 General life history 1919.4.1.4 Feeding 1919.4.1.5 Behaviour/adaptability 1919.4.1.6 Fishery status 1929.4.1.7 Aquaculture attributes and challenges 192

9.4.2 Broodstock management and hatchery operations 1939.4.2.1 Disease 194

9.5 Product Description 1949.6 Markets 1959.7 Future R & D for Shortnose and Atlantic Sturgeons 196

10 Milkfi sh (Family: Chanidae) 200Cheng-Sheng Lee and Chin-fa Liu

10.1 General Introduction 20010.2 Farming of Milkfish 20110.3 Broodstock Management and Hatchery Operations 20310.4 On-growing to Market Size 20810.5 Future Perspectives 212

11 The Catfi sh (Family: Ictaluridae) 216Jeffrey Silverstein, Terrill Hanson and Brian Bosworth

11.1 General Introduction 21611.2 Farming of Catfish 21711.3 Broodstock Management and Hatchery Operations 21911.4 On-growing to Market Size 22211.5 Future Perspectives 230

12 The Salmonids (Family: Salmonidae) 234Malcolm Jobling, Arne-Mikal Arnesen, Tillman Benfey, Chris Carter, Ronald Hardy, Nathalie R. Le François, Robyn O’Keefe, Juha Koskela and Simon G. Lamarre

12.1 General Introduction 23412.2 Whitefishes: Biology and Culture 236

12.2.1 Common whitefish, Coregonus lavaretus 23712.2.1.1 Farming of common whitefish 23912.2.1.2 Broodstock management and 239

hatchery operations11.2.1.3 On-growing to market size 240

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x Contents

12.3 Charrs: Biology and Culture 24112.3.1 Arctic charr, Salvelinus alpinus 242

12.3.1.1 Farming of Arctic charr 24512.3.1.2 Broodstock management and 245

hatchery operations12.3.1.3 On-growing to market size 247

12.3.2 Brook charr, Salvelinus fontinalis 24812.3.2.1 Farming of brook charr 25012.3.2.2 Broodstock management and 250

hatchery operations12.3.2.3 On-growing to market size 25212.3.2.4 Future perspectives 254

12.4 Atlantic Salmons and Trouts: Biology and Culture 25512.4.1 Atlantic salmon, Salmo salar 255

12.4.1.1 Life cycle of Atlantic salmon 25712.4.1.2 Farming of Atlantic salmon 26112.4.1.3 Broodstock management and 261

hatchery management12.4.1.4 Smolt production 26312.4.1.5 On-growing to market size 266

12.5 Pacific Salmons and Trouts: Biology and Culture 27112.5.1 Rainbow trout, Oncorhynchus mykiss 273

12.5.1.1 Life cycle of rainbow trout 27512.5.1.2 Farming of rainbow trout 27512.5.1.3 Broodstock management and 276

hatchery operations12.5.1.4 On-growing to market size 277

12.6 Acknowledgements 281

13 The Codfi shes (Family: Gadidae) 290Björn Björnsson, Matthew Litvak, Edward A. Trippel and Marc Suquet

13.1 General Introduction 29013.2 Atlantic Cod, Gadus morhua 290

13.2.1 Farming of Atlantic cod 29413.2.2 Rearing technologies and practices: full cycle 296

versus on-growing13.2.2.1 Capture-based cod farming 29713.2.2.2 Full-cycle cod farming 297

13.2.3 Broodstock management and 298hatchery operations

13.2.4 On-growing to market size 30013.2.5 Commercialization 30213.2.6 Future perspectives 303

13.3 Haddock, Melanogrammus aeglefinus 30413.3.1 Farming of haddock 307

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Contents xi

13.3.2 Broodstock management and 308hatchery operations

13.3.3 On-growing to market size 30913.3.4 Commercialization 30913.3.5 Future perspectives 310

13.4 Pollack, Pollachius pollachius 31013.4.1 Farming of pollack 31313.4.2 Broodstock management and 313

hatchery operations13.4.3 On-growing to market size 31313.4.4 Future perspectives 314

14 The Snooks (Family: Centropomidae) 323Chris Carter, Brett Glencross, Robin S. Katersky and Michel Bermudes

14.1 General Introduction 32314.2 Barramundi, Lates calcarifer 324

14.2.1 Farming of barramundi 32614.2.2 Broodstock management and husbandry 32714.2.3 On-growing to market size 32914.2.4 Commercialization 33214.2.5 Future perspectives 333

15 The Temperate Basses (Family: Moronidae) 337Malcolm Jobling, Stefano Peruzzi and Curry Woods

15.1 General Introduction 33715.2 European Seabass, Dicentrarchus labrax 338

15.2.1 Seabass biology 34115.2.2 Farming of seabass 34215.2.3 Broodstock management and 343

hatchery operations15.2.4 On-growing to market size 34715.2.5 Concluding comments 349

15.3 Striped Bass, Morone saxatilis, and Striped 350Bass Hybrids

15.3.1 Striped bass biology 35015.3.2 Farming striped bass and its hybrids 35115.3.3 Broodstock management and 351

hatchery operations15.3.4 On-growing to market size 35415.3.5 Concluding comments 356

16 Seabreams and Porgies (Family: Sparidae) 361Malcolm Jobling and Stefano Peruzzi

16.1 General Introduction 361

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xii Contents

16.2 Gilthead Seabream, Sparus aurata 36216.2.1 Farming of seabream 36516.2.2 Broodstock management and 367

hatchery operations16.2.3 On-growing to market size 37016.2.4 Concluding comments 372

17 Tilapia (Family: Cichlidae) 374Kevin Fitzsimmons and Wade O. Watanabe

17.1 General Introduction 37417.2 Tilapia, Oreochromis sp. 375

17.2.1 Broodstock management and 378hatchery operations

17.2.2 On-growing to market size 38117.2.3 Nutrition 38417.2.4 Impacts of introductions and farming 38617.2.5 Diseases 388

17.2.5.1 Viral 38817.2.5.2 Bacterial 38817.2.5.3 Mycotic 38817.2.5.4 Parasites 38917.2.5.5 Flagellates 38917.2.5.6 Ciliates 38917.2.5.7 Saltwater parasites 389

17.2.6 Production and markets for Tilapia 389

18 Drum-fi sh or Croakers (Family: Sciaenidae) 397Juan P. Lazo, Joan G. Holt, Christian Fauvel, Marc Suquet and Loic Quéméner

18.1 General Introduction 39718.2 Red Drum, Sciaenops ocellatus 398

18.2.1 Farming of red drum 40018.2.2 Broodstock management and 401

hatchery operations18.2.3 On-growing to market size 40218.2.4 Conclusion 405

18.3 Meagre, Argyrosomus regius 40518.3.1 Farming of meagre 40818.3.2 Broodstock management and 409

hatchery operations18.3.3 On-growing to market size 41018.3.4 Concluding comments 41218.3.5 Conclusion 412

19 The Wolffi shes (Family: Anarhichadidae) 417Nathalie R. Le François, Helge Tveiten, Laura C. Halfyard and Atle Foss

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Contents xiii

19.1 General Introduction 41719.2 Spotted Wolffish, Anarhichas minor, and 421

Atlantic Wolffish, A. lupus19.2.1 Farming of spotted wolffish 42219.2.2 Broodstock management and husbandry 42219.2.3 On-growing to market size 42519.2.4 Concluding comments 427

20 The Tunas (Family: Scombridae) 432Chris Carter, Barbara Nowak and Steven Clarke

20.1 General Introduction 43220.2 Farming of Tuna, Thunnus sp. 43420.3 Atlantic Bluefin Tuna, Thunnus thynnus 43520.4 Southern Bluefin Tuna, Thunnus maccoyii 43820.5 Pacific Bluefin Tuna, Thunnus orientalis 443

21 The Flatfi shes (Order: Pleuronectiformes) 450Albert K. Imsland

21.1 General Introduction 45021.2 Turbot, Scophthalmus maximus 454

21.2.1 Farming of turbot 45721.2.2 Broodstock management and 457

hatchery operations21.2.3 On-growing to market size 45921.2.4 Commercialization 46321.2.5 Future perspectives 463

21.3 Atlantic Halibut, Hippoglossus hippoglossus 46421.3.1 Farming of Atlantic halibut 46721.3.2 Broodstock management and 467

hatchery operations21.3.3 On-growing to market size 46921.3.4 Commercialization 47221.3.5 Future perspectives 472

21.4 Winter Flounder, Pseudopleuronectes americanus 47321.4.1 Farming of winter flounder 47321.4.2 Broodstock management and husbandry 47521.4.3 On-growing to market size 47621.4.4 Commercialization 47621.4.5 Future perspectives 477

21.5 Sole, Solea solea and Solea senegalensis 47821.5.1 Farming of sole 48121.5.2 Broodstock management and hatchery 482

operations (S. solea)21.5.3 Broodstock management and hatchery 483

operations (S. senegalensis)21.5.4 On-growing of S. solea to market size 48521.5.5 On-growing of S. senegalensis to market size 486

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xiv Contents

21.5.6 Commercialization 48721.5.7 Future perspectives 488

III Market and Economic Analysis 497

22 Marketing New Species 499James A. Young

22.1 Introduction 49922.2 What is Marketing? 50122.3 The Fish Marketing Environment 50322.4 Marketing Decisions on New Species 50622.5 Planning for New Species and Products 50822.6 Conclusion 511

23 Diversifi cation Pays: Economic Perspectives 514on Investment in Diversifi ed AquacultureJames R. Wilson and Bruno Archer

23.1 Introduction 51423.2 Economic Reasons for Diversification 51723.3 The Political Economics of Aquaculture Diversification 52023.4 A Case Study: Application to the Province of Québec 522

23.4.1 Private venture capital markets and diversified 523aquaculture

23.4.2 Public investment strategies 52423.5 Concluding Remarks 527

IV Future Perspectives 531

24 Offshore and Recirculation Technologies 533Richard Langan and Michel Couturier

24.1 Introduction 53324.2 Offshore Technologies 533

24.2.1 The context for offshore farming 53324.2.2 Current status of offshore finfish aquaculture 534

24.2.2.1 Defining offshore aquaculture 53424.2.2.2 Engineering design and 534

assessment24.2.2.3 Containment technologies 53524.2.2.4 Operational systems 53624.2.2.5 Species cultivated in 537

offshore cages24.2.2.6 Environmental considerations 538

24.2.3 Future prospects and challenges 53824.3 Recirculation Technologies 539

24.3.1 Recirculating aquaculture systems (RAS) 539

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Contents xv

24.3.2 Potential of RAS for species diversification 54024.3.3 Operation and design 54124.3.4 Future prospects and challenges 542

25 Valorization of Aquaculture By-products 546Pierre U. Blier and Nathalie R. Le François

25.1 Introduction 54625.2 Feed and Food Use of By-products 54825.3 Specialized Products 549

25.3.1 Enzymes 54925.3.2 Antifreeze proteins 55025.3.3 Antimicrobial polypeptides 551

25.4 A Case Study: Wolffish, Anarhichas lupus 551and A. minor

25.5 Concluding Remarks 553

26 Organic and Ecolabelling 556James A. Young

26.1 Introduction 55626.2 Production Pressures on Diversification 55726.3 The Role of Branding in Differentiation 55826.4 Organic Differentiation 56026.5 Market Impacts of Differentiation 56126.6 Ecolabelling 56226.7 Conclusion 564

27 The Future of Aquaculture: Insights from 567Economic TheoryGunnar Knapp

27.1 Introduction 56727.2 Supply and Demand Model 56727.3 Applying Supply and Demand Modelling to Aquaculture 57027.4 Modelling How Aquaculture May Change in the Future 57127.5 Technological Innovation in Aquaculture 57427.6 The Importance of Marketing for Aquaculture 57627.7 Effects of Government Policies on Aquaculture 57827.8 Modelling Diversification of Aquaculture 58227.9 Conclusions 584

Species factsheets: Appendix (1–24) 587Index 661

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xvii

Contributors

Bruno Archer, Centre Aquacole de la Côte-Nord (CACN), North-Shore Aquaculture Center (NSAC), 106 rue Napoléon, Suite 200-B, Sept-Îles (Québec), Canada G4R 3L7 ([email protected])

Arne-Mikal Arnesen, Nofima A/S, Muninbakken 9–13, Postboks 6122, N-9291 Tromsø, Norway ([email protected])

Tillman Benfey, Department of Biology, University of New Brunswick, PO Box 4400, Fredericton, New Brunswick, Canada E3B 6E1 ([email protected])

Michel Bermudes Government of Western Australia, Department of Fisheries, Research Division, PO Box 20, North Beach, WA 6920, Australia

Björn Björnsson, Marine Research Institute, Skulagata 4, 121 Reykjavik, Iceland ([email protected])

Pierre U. Blier, Département de biologie, chimie et géographie, Université du Québec à Rimouski, 300 Allée des Ursulines, Rimouski, QC, Canada G5L 3A1 ([email protected])

Brian Bosworth, US Department of Agriculture, Agricultural Research Service, Catfish Genetics Research Unit, Stoneville, USA

Chris Carter, University of Tasmania, Launceston Campus, Locked bag 1399, Tasmania 7250, Australia ([email protected])

Steven Clarke, Aquafin CRC, Box 120, Henley Beach, SA 5022, Australia ([email protected])

Michel Couturier, Department of Chemical Engineering, University of New Brunswick, PO Box 4400, Fredericton, NB, Canada E3B 5A3 ([email protected])

Lawrence Dooley, Department of Management, Centre for Wine, Food and Agribusiness, Location Room 03, Building N6, Caulfield, Monash University, Victoria 3800, Australia ([email protected])

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xviii Contributors

Christian Fauvel, Ifremer, Laboratoire LALR, Station expérimentale de Palavas, Chemin de Maguelone 34250 Palavas-les-Flôts, France ([email protected])

Kevin Fitzsimmons, Environmental Research Lab, University of Arizona, 2601 E Airport Dr, Tucson, AZ 85706-6985, USA ([email protected])

Atle Foss, Akvaplan-Niva A/S, Bergen office, PO Box 2026, Nordnes 5, N-5817, Bergen, Norway ([email protected])

Brett Glencross Government of Western Australia, Department of Fisheries, Research Division, PO Box 20, North Beach, WA 6920, Australia ([email protected])

Laura C. Halfyard, Marine Institute, Memorial University of Newfoundland, PO Box 4220, St John’s, Newfoundland, Canada A1C 5R3 ([email protected])

Terrill Hanson, Department of Fisheries and Allied Aquacultures, Auburn Univer-sity, 203 Swingle Hall, Auburn, AL 36849, USA ([email protected])

Ronald Hardy, UI Hagerman Fish Experiment Station, 3059 F National Fish Hatchery Road, Hagerman, Idaho 83332, USA ([email protected])

Joan G. Holt, University of Texas at Austin, Marine Science Institute, 750 Channel View Drive, Port Aransas, TX 78373-5015, USA ( [email protected])

Albert Imsland, Akvaplan-Niva A/S, Iceland Office, Akralind 4, 201 Kópavogi, Iceland and Department of Biology, University of Bergen, High Technology Centre, 5020 Bergen, Norway ([email protected])

Malcolm Jobling, University of Tromsø, 9037 Tromsø, Norway ([email protected])

Robin Katersky University of Tasmania, Locked bag 1370, Launceston, TAS 7250, Australia ([email protected])

Gunnar Knapp, Institute of Social and Economic Research, University of Alaska Anchorage, 3211 Providence Drive, Anchorage, Alaska 99508 ([email protected])

Juha Koskela, Finnish Game and Fisheries Research Institute, Laukaa Fisheries Research and Aquaculture, 41360 Valkola, Finland ( [email protected])

Simon G. Lamarre, Memorial University of Newfoundland, Canada ([email protected])

Richard Langan, Atlantic Marine Aquaculture Center, University of New Hampshire, Gregg Hall, 35 Colovos Road, Durham, NH 03824, USA ([email protected])

Juan P. Lazo, CICESE, Carretera Tijuana – Ensenada CP 22860, Ensenada, BC Mexico ([email protected])

Nathalie R. Le François, Biodôme de Montréal, 4777 Avenue Pierre-De Coubertin, Montréal, Québec H1V 1B3, Canada and Université du Québec à Rimouski, 300 des Ursulines, Rimouski, QC, G5L 3A1, Canada ([email protected])

Cheng-Sheng Lee, Center for Tropical and Subtropical Aquaculture Oceanic Institute, 41–202 Kalanianaole Highway, Waimanalo, HI 96795, USA ([email protected])

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Contributors xix

Matthew Litvak, Centre for Coastal Studies and Aquaculture and Department of Biology, University of New Brunswick, P.O. Box 5050, Saint John, New Brunswick, Canada E2L 4L5 ([email protected])

Chin-fa Liu, Consultant for Team Aqua Corporation, 14 Floor-1, No. 115, Jungshan Road, Shinying City, Tainan, Taiwan, 730, ROC ([email protected])

Barbara Nowak, School of Aquaculture, University of Tasmania, Launceston, Australia ([email protected])

Robyn O’Keefe, Department of Biology, University of New Brunswick, PO Box 4400, Fredericton, New Brunswick, Canada E3B 5A3 ([email protected])

David Otton, 23 Thompson Drive, Tathra, New South Wales, Australia 2550 ([email protected])

Stefano Peruzzi, University of Tromso, 9037 Tromsø, Norway ([email protected])

Loic Quéméner, Ifremer, Centre de Brest, BP70, 29280 Plouzané, France ([email protected])

Arianne Savoie, Université du Québec à Rimouski, 300 des Ursulines, Rimouski, QC, G5L 3A1, Canada ([email protected])

Jeffrey Silverstein, USDA/ARS National Center for Cool and Cold Water Aquaculture, 11861 Leetown Road, Kearneysville, WV 25430, USA ( [email protected])

Marc Suquet, Ifremer, Laboratoire ARN, Station Expérimentale d’Argenton, Presqu’île du Vivier, 29840 Landunvez, France ([email protected])

Edward A. Trippel, St Andrews Biological Station, 531 Brandy Cove, St Andrews, New Brunswick, Canada E5B 2L9 ([email protected])

Helge Tveiten, Nofima A/S, Muninbakken 9–13, Postboks 6122, N-9291 Tromsø, Norway ([email protected])

Wade O. Watanabe, Centre of Marine Science, University of North Carolina at Wilmington, 7205 Wrightsville Ave, Wilmington, NC 28403, USA ([email protected])

Guillaume Werstink, Université du Québec à Rimouski, 300 des Ursulines, Rimouski, QC, G5L 3A1, Canada ([email protected])

James R. Wilson, Département Économie et de Gestion, Université du Québec à Rimouski, 300 Allée des Ursulines, Rimouski, QC, Canada G5L 3A1 ([email protected])

Curry Woods, Department of Animal and Avian Sciences University of Maryland College Park, MD 20742, USA ([email protected])

James A. Young, Department of Marketing, University of Stirling, Stirling FK9 4LA, Scotland, UK ( [email protected])

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xxi

We would like to thank the following people:Brian Colwill and Dr David Otton for their assistance with linguistic verifica-

tion. David came to aquaculture as a mature aged PhD student and his single minded advocacy for the agribusiness approach and his enthusiasm for aquaculture will be greatly missed in Australia and beyond;

Marc Veillet and Karine Bisson of the Ministère de l’Agriculture, des Pêcheries et de l’Alimentation du Québec;

Dr Céline Audet (ISMER/UQAR, Québec, Canada) for reviewing the winter flounder section, Dr Denis Coves (Ifremer, France) for fruitful exchange on tuna cultivation efforts in Europe and Dr Serge Parent (Biodôme de Montréal, Québec, Canada) for his constructive comments;

François Larouche, Timothée Govarre and Sarah Tremblay-Bourgeois for punctual, highly specific tasks that provided relief when most needed.

The financial contributions of the Université du Québec à Rimouski and the Biodôme de Montréal are also acknowledged.

Acknowledgements

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I Aquaculture Diversification: an Introduction

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©CAB International 2010. Finfish Aquaculture Diversification(eds N.R. Le François, M. Jobling, C. Carter and P.U. Blier) 3

1 Fish Culture: Achievements and Challenges

MALCOLM JOBLING

University of Tromsø,Tromsø, Norway

1.1 Introduction

Aquaculture is the cultivation of aquatic organisms (e.g. algae, molluscs, crusta-ceans, fish) and implicit in this is human intervention that involves some control over the stock. The degree of human intervention may be limited in extensive culture of algae and molluscs and in extensive pond culture of fish and crusta-ceans, but the level of control exerted is high in intensive farming systems. Intensive culture involves intervention at all phases of the production cycle; from broodstock maintenance and egg production to the harvesting and marketing of the finished product (Fig. 1.1). The farmed animals are reared in captivity throughout their lives; they are held under relatively benign, semi-controlled conditions and rely on formulated feeds for their nutrition. Aquaculture research and development (R & D), as it relates to intensive fish farming, currently encom-passes all aspects of production and postharvest processing, including culture unit design, feed delivery systems, water treatment, feed formulation and dis-ease diagnosis and treatment. There is a growing perception that intensive aquaculture is a modern industry; this industry is becoming subject to increased scrutiny by policy makers and the general public and is becoming increasingly influenced by social attitudes that have an impact on regulations, marketing and product ranges and acceptability (Beardmore and Porter, 2003; Lee, 2003; Aerni, 2004; Kelso, 2004; Boyd et al., 2005; Focardi et al., 2005; Foran et al., 2005; Logar and Pollock, 2005; Midtlyng, 2005; Muir, 2005; Myhr and Dalmo, 2005; Naylor et al., 2005; Verbeke et al., 2005; Devlin et al., 2006; Huntingford et al., 2006).

Intensive fish culture is benefiting from increased use of sophisticated instrumentation and management tools, and modern analytical techniques are finding increased application in aquaculture R & D. In the sections that follow, examples will be given of the ways in which some of these recently developed

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4 M. Jobling

techniques have been used to address research problems with direct bearing on intensive aquaculture. There will also be consideration of some of the chal-lenges and contentious issues that the industry must face.

1.2 Application of Emerging Technologies to Aquaculture Research

In recent years, there have been major advances in a variety of analytical methods and molecular technologies. For example, molecular diagnostics can play a key role in the detection and identification of food contaminants and allergens, potential pathogens and genetically modified organisms (GMOs). Within this sphere, DNA microarray technology has created the possibility for simultaneous analysis of large gene sets, making it a potent molecular diagnos-tics tool with a number of applications (Liu-Stratton et al., 2004; Kato et al.,2005; Roy and Sen, 2006). As such, the development of molecular diagnostics techniques has had major impacts on R & D in a host of applied sciences, including clinical (human) and veterinary medicine, food science, agronomy and agricultural sciences.

These disciplines interface with aquaculture across a broad range of subject areas; genetics and selective breeding, health management and disease treat-ment, authentication and control of feed ingredients, postharvest processing, product storage and spoilage, assurance of food safety, product development and identification and traceability of products through the food chain (Blanco and Villarroya, 2002; Cozzolino et al., 2002, 2005a,b; Ali et al., 2004; Carbonaro, 2004; Dunham, 2004; Huss et al., 2004; Kuiper et al., 2004; Pérez-Marin

Fig. 1.1. Overview of the production cycle of farmed fish with an indication of the areas in which there is human intervention.

Animal improvement

Selective breeding

Sex reversal

Genetic biotechnologies

Broodstock

Gametes(eggs and sperm)

Larvae and juveniles

Grow-outto market size

Feedsand

feeding

Water and waste treatment

Water inflow Wastewater and effluent

Harvestingand

processing

Distributionand sales

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Achievements and Challenges 5

et al., 2004; Deisingh and Badrie, 2005; Focardi et al., 2005; Gorris, 2005; Guérard et al., 2005; Kelly et al., 2005; Lehrer and Bannon, 2005; Midtlyng, 2005; Monis et al., 2005; Neira and Diaz, 2005; Pinotti et al., 2005; Verbeke et al., 2005; Adams and Thompson, 2006; Berrini et al., 2006; Bruhn and Earl, 2006; Gasser, 2006; Reid et al., 2006; Toldrá and Reig, 2006). Similar sets of methods, such as spectroscopy (e.g. near-infrared (NIR), mid-infrared (MIR), UV-, Raman-, nuclear magnetic resonance (NMR) spectroscopy), immu-nological assays and DNA-based technologies, are being applied increasingly to assist in the solving of research problems in medical science, food science, agri-culture and livestock production and aquaculture.

Of the spectroscopic techniques, NIR spectroscopy has attained popularity because it is a rapid and non-destructive technique, equipment costs are rela-tively low and the equipment is quite easy to use. Thus, interest in the exploita-tion of the analytical potential of NIR paralleled the development of electronic optical equipment and computers that were capable of rapid extraction and processing of the information contained within NIR spectra. NIR spectroscopy can be used on both solid and liquid samples without any major pretreatment, and interpretation of the spectra can give information about both the physical and chemical properties of the sample (Blanco and Villarroya, 2002; Reid et al., 2006). It currently has use as a rapid analytical tool in quality control in the petrochemical and pharmaceutical industries and as an aid in diagnostics in the clinical and biomedical fields, but NIR spectroscopy has widest application in the food science and animal feed sectors. NIR spectroscopy is, for example, used for determining chemical composition (protein, moisture and oil/fat) of feed ingredients and feeds and of crops and livestock, either prior to or at har-vest/slaughter, and may also be used for identification and authentication pur-poses (Blanco and Villarroya, 2002; Cozzolino et al., 2002, 2005a,b; Pérez-Marin et al., 2004; Reid et al., 2006).

As a second example, the development of molecular marker techniques created new possibilities for the identification and genetic characterization of species, populations and individuals. This has important implications for the study of interrelationships between species (taxonomy and phylogeny), popula-tion genetics, selective breeding and genetics and disease diagnosis (Melamed et al., 2002; Ali et al., 2004; Dunham, 2004; Deisingh and Badrie, 2005; Gjedrem, 2005; Guérard et al., 2005; Monis et al., 2005; Neira and Diaz, 2005; Pinotti et al., 2005; Adams and Thompson, 2006; Chistiakov et al.,2006; Gasser, 2006). Methods involving the polymerase chain reaction (PCR) to amplify nucleic acids have proven particularly valuable and have been applied to several types of research problem. PCR, which has been in use for almost 20 years, is a fundamental technology for genetic identification and characteri-zation. DNA markers derived from PCR amplification are now being widely used for genetic characterization and for distinguishing between genetic vari-ants at different hierarchical levels: species, populations, strains and individuals (Ali et al., 2004; Dunham, 2004; Liu and Cordes, 2004; Gjedrem, 2005; Neira and Diaz, 2005; Chistiakov et al., 2006; Reid et al., 2006; Renshaw et al., 2006). Major advantages of PCR over alternative techniques for nucleic acid amplification are cost and ease of use. Using PCR, enzymatic amplification

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6 M. Jobling

of either mitochondrial or genomic DNA can be achieved in vitro from minute amounts of material. Highly conserved DNA regions can be used to delineate taxonomic relationships between species, examination of moderately con-served regions can be used to distinguish between populations, whereas varia-ble DNA regions can be used for tracking individuals and ‘mapping’ can be used to identify genetic markers that correlate genotype to phenotype, for example, linked to important production traits. The latter finds application in the identification of molecular markers of qualitative and quantitative trait loci (QTLs) of economic interest (Melamed et al., 2002; Ali et al., 2004; Dunham, 2004; Liu and Cordes, 2004; Neira and Diaz, 2005; Chistiakov et al., 2006). Thus, PCR encompasses amplification of a conserved region of mDNA for species identification as in PCR-RFLP, or a fingerprinting approach using PCR-RAPD for the detection of genetic variation or the definition of genetic markers.

1.3 Stock Improvement

Farmed stock can be improved by selective breeding and/or by implementing a variety of genetic biotechnologies (Maclean and Laight, 2000; Lee, 2001; Devlin and Nagahama, 2002; Melamed et al., 2002; Utter and Epifanio, 2002; Beardmore and Porter, 2003; Fjalestad et al., 2003; Dunham, 2004; Liu and Cordes, 2004; Gjedrem, 2005; Mignon-Grasteau et al., 2005; Neira and Diaz, 2005; Chistiakov et al., 2006; Goetz et al., 2006). DNA marker methods, and other genetic technologies, are having increased impact on the improvement of production characteristics of aquaculture stock and they find application in broodstock management and selective breeding programmes. For example, several fish species are currently under study for genome sequen-cing (Dunham, 2004; Volff, 2005; Goetz et al., 2006) and some commercially important species have been included in such investigations. These include channel catfish, Ictalurus punctatus, Atlantic salmon, Salmo salar, rainbow trout, Oncorhynchus mykiss, and Nile tilapia, Oreochromis niloticus (Fjalestad et al., 2003; Dunham, 2004; Volff, 2005), where the focus has been on the identification of QTLs of economic interest. Such QTLs relate to those with an influence on growth, age-at-maturation and reproductive characters, body form and pigmentation, environmental tolerances, disease resistance and behaviour (Fjalestad et al., 2003; Dunham, 2004; Liu and Cordes, 2004; Gjedrem, 2005; Neira and Diaz, 2005; Volff, 2005; Chistiakov et al., 2006), and identi-fied markers are being used to assist with broodstock selection for inclusion in breeding programmes.

Genetic interventions, environmental manipulations and hormonal treat-ments, either alone or in combination, may be used to influence sex determina-tion, sexual differentiation and reproductive physiology and behaviour (Lee, 2001; Patiño, 2002; Melamed et al., 2002; Kagawa et al., 2003; Dunham, 2004; Gjedrem, 2005; Neira and Diaz, 2005; Goetz et al., 2006; see Chapter 2, this volume). For example, photothermal manipulations are used to influence the onset of sexual maturation (timing of puberty) and are employed

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Achievements and Challenges 7

in broodstock management for production of out-of-season eggs. In addition, hormone treatments are used to induce gamete maturation and spawning in a wide range of species.

1.3.1 Sex-reversal and production of monosex populations and triploids

Genetic techniques, involving a sex-reversal step, can be used to produce single-sex (monosex) populations and gene manipulation may be employed to induce sterility through triploidization (Fig. 1.2). The production of monosex (single-sex) populations may be desirable when there are differences between the sexes in growth rate, age and size at maturity, disease resistance and other production characters, e.g. coloration, flesh characteristics, etc. The rearing of monosex populations may also be used to prevent unwanted (uncontrolled) spawning in rearing units such as ponds. The raising of sterile fish may be desirable as a measure to eliminate spawning in the farmed population and to reduce the potential negative genetic effects of farm escapees on natural popu-lations (due to interbreeding).

Production of sterile fish may be used to reduce the negative effects of sex-ual maturation on growth and prevent the deterioration of flesh quality (due to the mobilization of reserves from the muscle) that accompanies maturation in

Broodstock

Genotypic femalePhenotypic male

Phenotypic femaleGenotypic female

Broodstock

'Female'sperm

Normal (female)ova

Monosex(all-female)

triploids

Pressure or thermalshock shortly after

fertilization

Young juvenilesAndrogen treatment

at low dosage

Monosex(all-female)

diploids

Grow outDiploids harvested

and marketed beforethey mature

Genotypic femalefertilized eggs

Fig. 1.2. Schematic representation of sex-reversal techniques and the methods used to produce monosex (all female) diploid and triploid fish. In this scheme, it is assumed that the female is the homogametic (XX) sex and the male is the heterogametic (XY) sex. Sex reversal is induced by treating fertilized eggs, or very young juveniles, with male sex steroid hormones (androgens). Triploidy is induced by subjecting fertilized eggs to a temperature or a pressure shock.

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8 M. Jobling

many fish species. Muscle protein degradation involves three major proteolytic enzyme systems: the lysosomal cathepsins, the calpain–calpastatin (calpain–CAST) system and the ubiquitin–proteasome system (Salem et al., 2004, 2005a,b, 2006). Enzymes of these three systems are expressed differently in fertile diploid and sterile triploid rainbow trout, O. mykiss, with the cathepsins seemingly playing the major role in the proteolysis of muscle proteins seen in maturing fish. On the other hand, the calpain–CAST system appears to play an important role in the regulation of muscle protein turnover during growth and in the mobilization of reserves that occurs under a fast. This system, along with the cathepsins, is also involved in the degradation of muscle proteins that occurs post-mortem, thereby having an influence on muscle softening, textural changes and fillet ‘quality’ (Salem et al., 2004, 2005a,b, 2006).

Although methods used for producing monosex and/or sterile populations are now considered routine for several species, they do not have universal application due to the fact that several anomalous effects have been observed. Anomalies may arise, for example, if the eggs and hatchlings are exposed to waters of particular temperatures or with certain chemical characteristics (Devlin and Nagahama, 2002; Patiño, 2002; Kagawa et al., 2003; Miyamoto and Burger, 2003; Mills and Chichester, 2005; Goto-Kazeto et al., 2006; see Chapter 2, this volume). This is seen in intensive commercial production of European seabass, Dicentrarchus labrax, in which about 75% of the farmed fish usually develop as males; male-biased populations are undesirable because males grow more slowly and mature earlier than females. A substantial propor-tion of the male fish mature before reaching harvest size, so production of monosex female populations would be desirable. Although sex manipulation of seabass has been achieved by exposing fish to exogenous sex steroid hor-mones, efforts to produce female monosex seabass populations using crosses involving sex-reversed individuals have not met with success. A major stumbling block for the application of sex-reversal techniques to this species is that the mechanisms responsible for sex determination and differentiation are not known with certainty (Piferrer et al., 2005).

1.3.2 Induction of sterility

Chromosomal manipulation methods are useful for the production of triploid and tetraploid individuals, but they are not 100% successful. Consequently, the populations produced inevitably will contain a small percentage of fertile dip-loids. It is possible to induce sterility via the production of hybrids and by other methods. The biotechnological methods involve the production of individuals that express antagonists for a function needed for fertility, suppression of devel-opment and differentiation of germ cells, or suppression of the expression of genes required for the biosynthesis of sex steroid hormones (Maclean and Laight, 2000; Melamed et al., 2002; Kagawa et al., 2003; Weidinger et al.,2003; Dunham, 2004; Slanchev et al., 2005). For example, research into primordial germ cell (PGC) development and function has given promising results (Raz, 2003; Weidinger et al., 2003; Slanchev et al., 2005). PGCs are

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Achievements and Challenges 9

the cells that ultimately give rise to the gametes, but these cells do not have their source in the embryonic tissue that develops into the gonads. In other words, PGCs initially migrate from their source to the genital ridges of the developing embryo and there is then differentiation of the gonads. Over time, the PGCs differentiate into eggs or sperm in the ovary or testis, respectively (Raz, 2003). Work carried out on zebrafish, Danio rerio, provides evidence of a key role of PGCs in sex determination and gonad differentiation. Ablation of PGCs in embryonic zebrafish gave rise to sterile adults, all of which were males (as assessed using morphological and behavioural criteria, including courtship and mating with females) (Slanchev et al., 2005). This research may pave the way for similar studies on commercially farmed fish species, with the aim of producing sterile populations. Such populations might have better growth and production characteristics than fertile conspecifics and sterility could also reduce the potential impact of farm escapees on wild populations.

PGCs may be transplanted from one species to another and preliminary data provide evidence that this method can be used to develop surrogate brood-stock that produce xenogenic donor-derived offspring (Takeuchi et al., 2004). Further, the germ cells may retain a high degree of developmental plasticity even after the commencement of differentiation (Okutsu et al., 2006). For example, testicular germ cells taken from the testes of mature rainbow trout, O. mykiss,and transplanted into undifferentiated embryonic gonads were shown to have retained developmental plasticity. These cells were capable of displaying sexual bipotency. In other words, the testicular germ cells developed into spermatozoa in male embryonic recipients, but differentiated into eggs when the recipient embryonic rainbow trout were females (Okutsu et al., 2006). This demonstrated that the differentiation and development of the testicular germ cells was not autonomous and hard-wired, but could be controlled and directed by the envi-ronment to which the cells were exposed. In addition, the gametes, either sperm or eggs, derived from the transplanted testicular germ cells were fully functional and could be used for the production of viable offspring (Okutsu et al., 2006). These approaches to producing functional eggs and sperm offer a powerful tool for the study of gamete differentiation and development in fish and may also have potential for the rapid production of inbred strains that express a range of commercially desirable genetic characters.

1.3.3 Interactions between farmed and wild fish

Although the production of sterile farmed stock might eliminate problems related to the genetic impacts of farm escapees on natural populations, it would not serve to eliminate all forms of interaction. Individuals that escape from farms could be members of non-native or introduced species, highly selected domesti-cated stocks, hybrids or animals that have had their genetic constitutions altered using transgenic biotechnologies (genetically modified organisms – GMOs). These different categories of exotics have different characteristics and have the potential to impact wild populations in several ways. The farm escapees could act as either predators or prey, they could be vectors of parasites or disease

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10 M. Jobling

organisms and they could compete for food or other habitat resources, resulting in displacement or exclusion of native species and natural populations. In the case of the farm escapees being fertile, there could also be interbreeding or hybridization between the exotics and native stocks, resulting in changes to genetic constitution or sterility. In all likelihood, several of these mechanisms would act simultaneously (Beardmore and Porter, 2003; Aerni, 2004; Bessey et al., 2004; Devlin et al., 2004, 2006; Kelso, 2004; Muir, 2004; Welcomme and Craig, 2004; Bondad-Reantaso et al., 2005; Boyd et al., 2005; Logar and Pollock, 2005; Murray and Peeler, 2005; Naylor et al., 2005; Bekkevold et al.,2006; Jonsson and Jonsson, 2006; Wessel et al., 2006).

1.3.4 Transgenic fish

Hitherto, over 30 species of fish have been the subject of transgenic research and genetic engineering, and the species studied include several representa-tives from the major farmed groups of fishes – the carps, tilapias, catfishes and salmonids. Transgenesis may be used to obtain basic information on gene func-tion and regulation, but also has commercial application in the improvement of animal production and in the introduction of gene constructs that result in the biosynthesis of high-value products such as recombinant proteins (Beardmore and Porter, 2003; Houdebine, 2005). The application of transgenic technol-ogy in fish has been concerned mostly with the examination of methods that can be used to enhance the growth of farmed stock, predominantly via the transfer of genes coding for growth hormone (Maclean and Laight, 2000; Beardmore and Porter, 2003; Dunham, 2004; Muir, 2004; Houdebine, 2005; Logar and Pollock, 2005; Devlin et al., 2006), but additional possibilities exist for exploitation of this type of genetic manipulation. For example, gene con-structs may be introduced to enhance disease resistance or modify metabolic pathways that have influences on the chemical composition of the flesh (Beardmore and Porter, 2003; Devlin et al., 2006).

One such example is the modification of fatty acid metabolic pathways to enhance the biosynthesis of n-3 highly unsaturated fatty acids (n-3 HUFAs) and increase their deposition in the flesh of the farmed animal (Alimuddin et al.,2005). The biosynthesis of HUFAs depends on a series of fatty acid desatura-tion and elongation reactions that are catalysed by a number of enzymes (Fig. 1.3). Fish species differ in their abilities to carry out the desaturation and elonga-tion reactions and the characteristics of the enzymes differ among species (Zheng et al., 2004; Agaba et al., 2005; Robert, 2006). Most notably, there are differences between carnivorous marine species and herbivorous and omniv-orous freshwater fish species in the ability to synthesize HUFAs from the 18C precursor fatty acids. Genes coding for key enzymes in the n-3 HUFA biosyn-thetic pathway may be introduced into the fish genome and fatty acid metabolic pathways modified accordingly (Alimuddin et al., 2005). There is, however, the possibility that introduction of novel DNA into the fish genome could give rise to unintended side effects, such as overexpression of some genes and the silenc-ing of others; the examination of unintended side effects of genetic engineering in fish is in its infancy, although metabolomics and other ‘omics-based’ methods

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Achievements and Challenges 11

for carrying out such investigations are available (Beardmore and Porter, 2003; Kuiper et al., 2004; Liu-Stratton et al., 2004; Kato et al., 2005; Griffin, 2006; Rischer and Oksman-Caldentey, 2006; Viant et al., 2006).

1.4 Feed Ingredients and Resources

Developments in plant breeding and genetics are playing an increasingly impor-tant role in the provision of ingredients for aqua-feeds. Although a wide range of ingredients is used in the formulation of aqua-feeds (Hertrampf and Piedad-Pascual, 2000), fishmeals and fish oils traditionally have been used as

Fatty acid series Enzyme

16:0 18:0

∆9 desaturase ↓ ↓

∆6 desaturase ↓ ↓ ↓ ↓

elongase ↓ ↓ ↓ ↓

∆5 desaturase ↓ ↓ ↓ ↓

elongase ↓ ↓ ↓ ↓

elongase ↓ ↓

∆6 desaturase ↓ ↓

(β-Oxidation) ↓ ↓

n-7 n-9 n-6 n-3

16:1 n-7 18:1 n-9 18:2 n-6 18:3 n-3

16:2 n-7 18:2 n-9 18:3 n-6 18:4 n-3

18:2 n-7 20:2 n-9 20:3 n-6 20:4 n-3

18:3 n-7 20:3 n-9 20:4 n-6 20:5 n-3

20:3 n-7 22:3 n-9 22:4 n-6 22:5 n-3

24:4 n-6 24:5 n-3

24:5 n-6 24:6 n-3

22:5 n-6 22:6 n-3

Fig. 1.3. Pathways for chain elongation and desaturation of fatty acids. Fatty acids of the n-7 and n-9 series can be synthesized de novo, whereas fatty acids of the n-6 and n-3 series are the essential fatty acids. The ∆6 desaturase enzyme has a preference for unsaturated fatty acids as substrate. In the absence of 18:3 n-3 and 18:2 n-6, 18:1 n-9 is desaturated and elongated, leading to an accumulation of long-chain fatty acids of the n-9 series in tissues (this can be used as an indication of essential fatty acid deficiency). Carnivorous marine fish species seem to have low ∆5 desaturase activity and they have a low capacity to use the n-3 and n-6 18C fatty acids as precursors for synthesis of n-3 and n-6 HUFAs. The shorthand notations of the fatty acids give information about the number of carbon atoms, the number of double bonds and the position of the first double bond from the methyl end of the molecule (from Jobling, 2004c).

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12 M. Jobling

major ingredients, but in recent years there has been a trend towards an increased reliance on plant-based products (see Chapter 3, this volume). Fishmeals have been used as a major ingredient because they have been a widely available source of protein of high quality. Marine fish oils have been widely used as the main lipid source in aqua-feeds because they have been readily available, relatively cheap and are an excellent source of the n-3 HUFAs deemed beneficial for human health (Graham et al., 2004; Ruxton et al.,2004, 2005; SACN-FSA, 2004; Bergé and Barnathan, 2005; Bourre, 2005; Cleland et al., 2005; MacRae et al., 2005; Napier and Sayanova, 2005; Bethune et al., 2006; Kim and Mendis, 2006). The heavy reliance on fish-meals and oils as major ingredients in aqua-feeds has, however, been ques-tioned and criticized (Naylor et al., 2000; Muir, 2005). Stocks of small, pelagic fish used for fishmeal and oil production represent finite resources that are either fully exploited or are being overexploited and are at risk of collapse. Fears have also been expressed that fishmeals and oils may give rise to con-tamination of the flesh of farmed fish with organochlorine compounds and/or heavy metals (Easton et al., 2002; Jacobs et al., 2002; Hites et al., 2004; Bell et al., 2005; Berntssen et al., 2005; Carlson and Hites, 2005; Foran et al.,2005; Hamilton et al., 2005; Bethune et al., 2006; Montory and Barra, 2006). These contaminants may be carcinogenic and may give rise to develop-mental anomalies that result in the malformation of various tissues and organs in both fish and human consumers (Miyamoto and Berger, 2003; Sumpter and Johnson, 2005; Scott et al., 2006; see Chapter 3, this volume).

1.4.1 Feed ingredients of plant origin

There may be major benefits to be gained by including protein sources of plant origin in fish feeds, but there are also problems associated with such feed sources (Hertrampf and Piedad-Pascual, 2000; McKevith, 2005). For exam-ple, many plants are deficient in some essential nutrients, such as some amino acids (e.g. lysine and methionine). This means that there may be a need to add amino acid supplements to feeds prepared with protein sources derived from plants. In addition, most plants contain antinutritional factors (ANFs) that inter-fere with digestion and absorption, reduce nutrient bioavailability and feed uti-lization and may have other adverse effects (Table 1.1) (Francis et al., 2001; Jobling, 2004c; Acamovic and Brooker, 2005). Further, it is becoming increas-ingly difficult to obtain vegetable feed ingredients, for example, soya, maize and canola, guaranteed to be free from some form of biotechnological genetic modification, i.e. non-transgenic vegetable feed ingredients are becoming increasingly scarce in the global market (Kok and Kuiper, 2003; Flachowsky et al., 2005; Myhr and Dalmo, 2005; Bruhn and Earl, 2006).

1.4.1.1 Transgenic plants

The genetic modification of plants may involve the introduction of genes encod-ing for increased disease resistance, production of insecticidal proteins or

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Table 1.1. Overview of major classes of antinutritional factors (ANFs) present in plants used as protein and/or carbohydrate sources in feedsprepared for farmed fish. × indicates that the ANF is present and heat-labile ANFs (i.e. ANFs that can be destroyed or denatured by heat treatment) are shown in bold typeface (from Jobling, 2004c).

Antinutritional factor (ANF)

Enzyme inhibitors

Protease Amylase Phytate Phyto-oestrogens Saponins Tannins Lectins Alkaloids Cyanogens Glucosinolates

‘Oilseed’ meals Soybean ¥ × × × ¥ Rape/canola ¥ × × × × Cottonseed ¥ × × Sunflower ¥ × × Sesame ¥Legumes Lupin ¥ × × × Lucerne × × × Faba bean ¥ × × ¥‘Grains’ Maize ¥ × × Wheat ¥ ¥ × × ¥ Sorghum ¥ ¥ × × ¥ בTubers’ Potato ¥ ¥ × ¥ × Cassava ¥ × ×

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increased herbicide tolerance; the agents that impart these properties are present at very low concentration and have little influence on the gross compo-sition of the plant. Alternatively, attempts may be made to influence nutritional value by increasing the protein content or indispensable (essential) amino acids, increasing concentrations of vitamins, such as tocopherols, or by reducing con-centrations of ANFs (Tucker, 2003; Flachowsky et al., 2005; Lehrer and Bannon, 2005; Bruhn and Earl, 2006; Dalal et al., 2006).

Plant oils are generally deficient in n-3 HUFAs, although several contain relatively high concentrations of 18C n-3 precursors (Hertrampf and Piedad-Pascual, 2000; McKevith, 2005; Robert, 2006). The fatty acid compositions of the flesh of monogastric animals frequently reflect the compositions of the feed, so aquaculture species given feeds based on plant oils may have rela-tively low concentrations of n-3 HUFAs in their flesh (e.g. Jobling, 2004a,b,c; Wonnacott et al., 2004; Berntssen et al., 2005; Torstensen et al., 2005; Visentainer et al., 2005; see Chapter 3, this volume). Given this, one major area of research focus has been the elucidation and modification of the meta-bolic pathways of fatty acid synthesis, with the aim of producing plants with higher n-3 HUFA content. The conventional metabolic pathway for the syn-thesis of the HUFAs is via the n-3 pathway employing the ∆6 desaturase and elongase route (Fig. 1.3), but there are alternative pathways. One of the alter-natives involves the n-6 pathway and a ∆17 desaturase (C20:4 n-6 → 20:5 n-3), another involves the n-3 pathway and a ∆4 desaturase (C22:5 n-3 →22:6 n-3) and there is also a route that relies on the n-3 pathway and ∆9elongase and ∆8 desaturase (C18:3 n-3 → 20:3 n-3 → 20:4 n-3). The alter-native pathways, found in some unicellular organisms, appear to offer most promise with regard to introduction into higher plants (Abbadi et al., 2004; Graham et al., 2004; Domergue et al., 2005; Flachowsky et al., 2005; Napier and Sayanova, 2005; Guschina and Harwood, 2006; Napier et al., 2006; Robert, 2006).

Animal feeds, including aqua-feeds, may also be supplemented with n-3 HUFAs obtained by microbial fermentation (Ward and Singh, 2005; Spolaore et al., 2006). The best microbial sources of these fatty acids are unicellular algae, or algal-like protists, of the order Thraustochytriales. Industrial fermenta-tion methods have been developed for several species and/or strains of Thraustochytrium and Schizochytrium (Bergé and Barnathan, 2005; Ward and Singh, 2005). Although plants can be engineered to produce n-3 HUFAs, microorganisms have certain advantages with regard to production, extraction and purification of the fatty acids. Several fatty acid products derived from fer-mentation of microorganisms are currently available commercially and are being incorporated into some aqua-feeds. The microorganism fatty acids are being used particularly in feeds destined for the rearing of the youngest devel-opmental stages of fish and crustaceans (Bergé and Barnathan, 2005; Ward and Singh, 2005; Guschina and Harwood, 2006; Spolaore et al., 2006).

Genetic modification of plants and animals by transgenesis gives rise to the possibility of introducing unintentional side effects as a result of the insertion of novel DNA into the genome. Possible results could be the increased biosynthesis and accumulation of toxic metabolites, enhanced production of known or novel

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allergenic proteins or reductions in tissue concentrations of certain essential nutrients in the modified organism (Kok and Kuiper, 2003; Kuiper et al., 2004; Liu-Stratton et al., 2004; Lehrer and Bannon, 2005; Rischer and Oksman-Caldentey, 2006; Roy and Sen, 2006). Such changes could have negative effects on an animal that consumes such products. Control measures have been introduced in an attempt to reduce such risks and a number of methods, includ-ing metabolic profiling or metabolomics (the global mapping of metabolites present within cells or tissues), are being developed to address these problems (Kuiper et al., 2004; Liu-Stratton et al., 2004; Kato et al., 2005; Lehrer and Bannon, 2005; Griffin, 2006; Rischer and Oksman-Caldentey, 2006; Müller and Steinhart, 2007). Some of these methods have been applied to examina-tion of the metabolic profiles of fish. These novel, modern methods have, how-ever, been used mostly in toxicological studies to identify changes in metabolism that result from the exposure of fish to toxicants or other forms of environmen-tal stressor (e.g. Cakmak et al., 2006; Viant et al., 2006).

The assessment of the effects of including genetically modified feed ingre-dients in fish feeds is in its infancy and has mostly involved examination of growth effects and histopathology (Brown et al., 2003; Glencross et al., 2003; Hemre et al., 2005; Sanden et al., 2005, 2006); hitherto, no major negative effects have been reported. Studies are also being conducted to examine the degree to which dietary DNA is degraded in the gut and to what extent it is taken up and distributed in body tissues; such studies are needed for assessment of the potential for horizontal gene transfer from genetically modified feed ingredients to the animal that consumes them (Sanden et al., 2004; Nielsen et al., 2005, 2006).

1.5 Health Management and Disease Control

Infectious diseases are a major problem in all intensive animal production sys-tems and intensive aquaculture is no exception; farm populations of aquatic animals may be decimated by a disease outbreak, leading to serious losses of production. Outbreaks of disease often occur when rearing conditions and ani-mal welfare are poor, so a suboptimal environment is often a major contribu-tory factor to such outbreaks. As such, health management starts with the creation of a suitable rearing environment and continues with the maintenance of that environment, including the minimizing of potential stressors (Håstein et al., 2005; Keeling, 2005; Midtlyng, 2005; Murray and Peeler, 2005; Adams and Thompson, 2006; Bricknell et al., 2006; Broom, 2006; Dawkins, 2006; Huntingford et al., 2006; Lund, 2006; see Chapter 2, this volume).

The stock should be provided with water of good quality, sufficient space and sufficient food of high nutritional quality and should also be subjected to limited disturbance and physical handling. The difference between health and disease depends on the balance between the pathogen and the host, and that balance can be influenced greatly by environmental factors such as water temperature and water quality (Bondad-Reantaso et al., 2005; Håstein et al.,2005; Murray and Peeler, 2005; Huntingford et al., 2006). Biosecurity has

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disease prevention and prophylaxis as its key points. It involves implementation of measures for prevention, control and eradication of economically important diseases. In other words, biosecurity encompasses all procedures required to protect the fish from contracting, carrying and spreading diseases and parasites (Bondad-Reantaso et al., 2005; Midtlyng, 2005; Murray and Peeler, 2005).

A major source of disease-causing organisms for aquatic animals is the water flowing into the rearing units, so treatment of incoming water is often used to reduce the levels of waterborne pathogens to below infective limits (see Chapter 2, this volume). Disinfection, which must inactivate a wide range of bacteria, viruses and protozoans, may be accomplished by chemical or physical means, such as chlorination, ozonation or treatment with UV or ionizing radia-tion. UV radiation treatment is frequently the method of choice for the disinfec-tion of water to be used for aquaculture purposes (Lee, 2003; Sharrer et al.,2005; Hijnen et al., 2006). An alternative, and not mutually exclusive alterna-tive, to disinfection to combat disease-causing organisms involves the use of probiotics. Probiotics may be defined as microorganisms that are administered to the fish, either via the water or in the feed, and they then enter the gastroin-testinal tract and serve a function in improving the health of the farmed stock (Gatesoupe, 1999; Irianto and Austin, 2002; Balcázar et al., 2006; Vine et al., 2006). The probiotics may act by stimulating the immune system of the host fish, or they may have direct effects on potential pathogens. The direct effects include interference with the metabolism of the pathogens via the pro-duction of inhibitory compounds by the probiotic microorganism and competi-tion for nutrients or other resources (Gatesoupe, 1999; Irianto and Austin, 2002; Balcázar et al., 2006; Vine et al., 2006).

1.5.1 Diagnostics and disease treatment

Veterinary medicine has long traditions in health surveillance and disease diag-nosis of terrestrial livestock, but incomplete knowledge about the environmen-tal requirements and biology of farmed aquatic animals along with a relative dearth of information about their diseases has meant that less has been achieved to solve the medical problems faced by aquaculture stocks (Woo et al., 2002; Storey, 2005; Bricknell et al., 2006). Molecular markers are being used for diagnostic and epidemiological purposes in clinical and veterinary medicine (Liu-Stratton et al., 2004; Monis et al., 2005; Adams and Thompson, 2006; Chistiakov et al., 2006; Gasser, 2006) and DNA-based technologies are find-ing increased use in the production of vaccines and pharmaceutical agents with human and animal health applications (Melamed et al., 2002; Houdebine, 2005; Midtlyng, 2005; Myhr and Dalmo, 2005; Dalal et al., 2006). As the aquaculture industry continues to grow, these areas will probably increase in priority as R & D objectives.

The development of diagnostic methods and vaccines against some of the diseases that attack aquaculture species has been instrumental in preventing disease outbreaks, thereby reducing the large-scale use of antibiotics and other drugs for the treatment of certain diseases. Nevertheless, there are relatively

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few commercial vaccines currently available and the pharmaceutical toolkit of the aquaculture veterinarian is also limited (Midtlyng, 2005; Sommerset et al.,2005; Storey, 2005; Bricknell et al., 2006; MacMillan et al., 2006).

1.5.1.1 Vaccines

Most vaccines are based on inactivated bacterial pathogens and few vaccines are available for other disease organisms, such as viruses (Midtlyng, 2005; Rigos and Troisi, 2005; Sommerset et al., 2005), although DNA vaccines against some viral agents are under development (Melamed et al., 2002; Midtlyng, 2005; Myhr and Dalmo, 2005; Sommerset et al., 2005; Adams and Thompson, 2006). DNA vaccines are gene constructs that express cloned antigens tran-siently in the treated individual, thereby promoting the development of protec-tion against the disease agent; R & D into DNA vaccines includes work on viral haemorrhagic septicaemia, infectious haematopoetic necrosis, channel catfish herpes virus, infectious salmon anaemia and infectious pancreatic necrosis.

Vaccines are usually administered by injection or by bath (immersion) treatment. The former is labour-intensive and time-consuming if large num-bers of animals are to be treated and bath treatments are often of reduced efficacy compared with injection, so alternative methods of vaccine delivery are being sought and developed (Vandenberg, 2004; Midtlyng, 2005; Adams and Thompson, 2006). Oral administration of vaccine via the feed offers the possibility of treating large numbers of animals with relative ease and avoids the trauma associated with handling and the injection of vaccines, but the efficacy of oral vaccination has, hitherto, proven variable (e.g. Esteve-Gassent et al., 2004; Romalde et al., 2004; Vandenberg, 2004; Shoemaker et al.,2006). Problems with oral vaccination relate to the possible degradation of vaccine components in the digestive tract, requiring the development of micro-encapsulation, and other methods, for vaccine protection.

1.5.1.2 Antibiotics and other chemotherapeutic agents

At present, the vaccines available target diseases prevalent in a limited number of farmed species, so the use of other disease treatments, such as antibiotics, is widespread in the aquaculture industry (Rigos and Troisi, 2005; Bricknell et al., 2006). Several antimicrobial agents, including tetracyclines, quinolones, sulphonamides and amphenicols, may be efficacious for the treatment of bacte-rial diseases of aquatic animals, but the range of drugs, pharmaceutical and other chemical agents approved for use in aquatic species is small compared with the number of drugs licensed for use in terrestrial livestock (Storey, 2005; MacMillan et al., 2006).

One major reason for this is that the size of the aquaculture market is cur-rently too small to offer drug companies an adequate return on the economic investment required to collect the documentation needed for approval to be granted by licensing authorities; the requirements may include documentation that the drug treatment is effective against the target disease in each specified aquatic animal, that the drug is safe for the treated animal, does not pose a hazard to humans who may consume the animal and does not give rise to

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environmental concerns (Midtlyng, 2005; Storey, 2005; MacMillan et al.,2006). These requirements impose difficulties for the development and imple-mentation of effective health management programmes for species that are cultured in low volume or that are under consideration for commercial farming (Storey, 2005; Bricknell et al., 2006). As the aquaculture industry expands, the data needed for the licensing of pharmaceuticals are gradually being col-lected and are being made available to private companies and licensing authori-ties (Bondad-Reantaso et al., 2005; Reimschuessel et al., 2005; Rigos and Troisi, 2005; Samuelsen, 2006).

Prophylactic use of vaccines and probiotics, combined with good manage-ment practices, reduces the incidence of disease outbreaks (Midtlyng, 2005; Sommerset et al., 2005; Balcázar et al., 2006; Vine et al., 2006), which, in turn, reduces the need to use antimicrobial drugs and chemotherapeutic agents (Murray and Peeler, 2005; Rigos and Troisi, 2005). Reduced reliance on such drug treatments has several benefits. For example, the treatment of water in reuse culture systems usually relies on biological action to remove metabolic wastes and the organisms responsible for the biodegradation of these wastes will be susceptible to some of the agents used to combat disease organisms. In this respect, the use of antibiotics, drugs and other chemical agents that kill water-treatment microorganisms would act as a double-edged sword.

Further, there is public concern regarding the use of drugs and other chemicals such as disinfectants, pesticides and chemotherapeutic agents in aquaculture. The general public is concerned about possible environmental contamination, and the negative effects that could result, and the possibility that development of antibiotic resistance in aquatic microorganisms could be transferred to human pathogens (Bondad-Reantaso et al., 2005; Focardi et al., 2005; Rigos and Troisi, 2005; Adams and Thompson, 2006; MacMillan et al., 2006; Toldrá and Reig, 2006). To help allay these concerns, the use of medicated feeds should be kept to a minimum, every effort should be made to prevent the loss of such feeds to the environment and all drugs, disinfectants and other chemical agents must be used in a responsible manner (Boyd et al.,2005; Focardi et al., 2005; Murray and Peeler, 2005; Rigos and Troisi, 2005; Storey, 2005; MacMillan et al., 2006; Toldrá and Reig, 2006).

1.6 Food Products: Nutritional Composition, Food Safety and Traceability

Aquaculture products intended as food should meet the tastes and expectations of the consumer, but there are many factors that interact to influence the com-position of these products. Most of these factors exert their influence prior to harvest, but postharvest processing, transport and storage conditions can also have a major influence. For example, the nature and nutrient composition of the feed are directly influential in determining flesh characteristics, but other agents, such as chemical therapeutic agents, drug residues and waterborne pollutants and contaminants that impart off-flavours and taints to the flesh, may also have direct effects (Fig. 1.4) (e.g. Tucker, 2000; Jobling, 2004a,b,c;

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Bell et al., 2005; Berntssen et al., 2005; Carlson and Hites, 2005; Torstensen et al., 2005; Bethune et al., 2006; Toldrá and Reig, 2006; see Chapters 2 and 3, this volume).

This means that attention must be given to food quality and safety at all links in the food supply chain from the producer to the consumer. The estab-lishment of clear objectives and accreditation procedures is integral to food quality and safety management. Increased control over the safety and quality of food is made possible by adopting agreed practices and implementing one or more management systems, such as good manufacturing practice (GMP), good hygiene practice (GHP) and hazard analysis critical control points (HACCP) (Huss et al., 2004; Sumner et al., 2004; Boyd et al., 2005; Gorris, 2005; da Cruz et al., 2006).

1.6.1 Analysis of aquaculture products

The versatility of NIR spectroscopy, and its advantages of speed and simplicity of instrument operation, has resulted in its adoption as a rapid analytical tool in many areas (Blanco and Villarroya, 2002; Reid et al., 2006), including the seafood and aquaculture industries. The main application of NIR spectroscopy within the seafood and aquaculture sectors has been for the prediction of

Feeds and feedingFeed composition (e.g. % fat, pigments)Feed contaminants (e.g. aflatoxins, dioxins)Feed amount and feeding frequency

Culture environment

Pollutants and contaminants (‘off-flavours’ and taints)Temperature, salinity and light conditionsWater quality (e.g. oxygen, metabolic wastes)Disease agents and parasites

Harvesting, processing,transport and storage

Species and stock (genetic selection)Life cycle stage, sex and maturityHealth status (diseases and parasites)

Fish characteristics

Fig. 1.4. Summary of some of the major factors influencing flesh (fillet) composition of farmed fish. The factors that play a role in determining the composition and ‘quality’ of the fish that reach the market are the intrinsic (relating to the fish themselves) and extrinsic factors (relating to feeds and feeding and the culture environment) that exert an influence during the growth cycle and the postharvest changes that occur during processing and fish storage.

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chemical (nutritional) composition, but more recent applications include spe-cies identification and authentication, monitoring of degradation products for assessment of freshness, spoilage and ‘quality’ and for discrimination between fresh and frozen-thawed fish (Cozzolino et al., 2002, 2005,a,b; Nilsen et al.,2002; Uddin and Okazaki, 2004; Nilsen and Esaiassen, 2005; Uddin et al.,2005; Lin et al., 2006; Reid et al., 2006).

Alternatives to NIR spectroscopy are available for the assessment of spoil-age via analysis of volatile compounds (e.g. aldehydes, ketones, organic acids and their esters and ethers). The composition and concentration of volatile compounds differs between fresh, stale and spoiled fish, with spoilage volatiles developing as a result of oxidative or microbial degradation of the chemical constituents of the tissues. Thus, as fish spoilage progresses, there is increased production of basic volatile nitrogen compounds, such as trimethylamine, ammonia and dimethylamine, and it is possible to detect changes in the con-centrations of these spoilage compounds remotely (Pacquit et al., 2006). There are also changes in other compounds as fish become spoiled, so there is a change in the odour profile (aromagram) over time and these changes can be used in quality assessment (Olafsdottir et al., 2004; Reid et al., 2006; Plutowska and Wardencki, 2007).

Other methods for identifying the species and/or sources of seafood, including aquaculture, products rely on molecular diagnostic techniques that involve the detection of some form of biomarker. Several methods, such as isoelectric focusing, electrophoresis and immunoassay, rely on the identifica-tion of specific proteins or groups of proteins for discrimination between spe-cies (Berrini et al., 2006), for identification of potential allergens (Lehrer et al.,2003; Van Do et al., 2005; Kobayashi et al., 2006; Müller and Steinhart, 2007), or for detection of therapeutic agents and drug residues that may be present in the products (Toldrá and Reig, 2006). For example, the major aller-gens present in seafood are muscle proteins – parvalbumins in fish and tropo-mysins in crustaceans – and immunoassays have been developed to detect the allergens present in a range of seafood species (Lehrer et al., 2003; Van Do et al., 2005; Kobayashi et al., 2006).

Other approaches to the rapid screening of food products for the assurance of quality and safety are based on the use of DNA microarrays (Liu-Stratton et al., 2004; Kato et al., 2005; Roy and Sen, 2006) and the development of electronic sensors, biosensors or ‘biochips’ (Toldrá and Reig, 2006; Müller and Steinhart, 2007). A biosensor consists of a chemical recogni-tion element or biological sensor, usually an antibody or enzyme, which is con-nected to a data processing unit. The target chemicals contact and bind to specific biological sensors, the biomolecular interaction signals are converted into electrical signals and the electrical signals are then processed by a micro-processor. Biochip array biosensors have the advantages that the method is rapid and that food products can be screened for several chemicals simultane-ously (as many as the number of specific recognition elements in the array), but initial investment costs for the development of biosensors are quite high.

A variety of other biomarker-based methods has also been developed for discrimination, authentication and quality evaluation purposes (Carbonaro,

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2004; Bergé and Barnathan, 2005; Guérard et al., 2005; Kelly et al., 2005; Pinotti et al., 2005; Reid et al., 2006; Renshaw et al., 2006; Toldrá and Reig, 2006; Müller and Steinhart, 2007). For example, techniques, such as PCR-RFLP and PCR-RAPD, which rely on DNA markers, have advantages over protein-based methods as they are less influenced by the tissue analysed, the age of the individual and the effects of thermal degradation. Using DNA-marker techniques, closely related species can be distinguished from each other and these techniques may enable recognition of the species compositions of com-plex mixtures and processed products (Chistiakov et al., 2006). An additional example is the use of fatty acid profiles for the discrimination of farmed from wild fish; the fatty acids can be used as biomarkers because the fatty acid pro-files of the animals generally reflect those of their food and the fatty acids present in aqua-feeds will usually differ from those present in the natural prey of the farmed species (e.g. Jobling, 2004a,b,c; Wonnacott et al., 2004; Bell et al., 2005; Bergé and Barnathan, 2005; Berntssen et al., 2005; Torstensen et al., 2005; Visentainer et al., 2005; see Chapter 3, this volume).

1.7 Fish as Food: Consumer Attitudes (and the Question of GM-Fish)

Price, availability, taste, nutritional value, attitudes and moral considerations all influence consumer food choice and there is increasing interest in food-related issues among the public at large (Shanahan et al., 2001; Butterworth, 2004; Huss et al., 2004; Bennett et al., 2005; Deisingh and Badrie, 2005; Frewer et al., 2005; Gorris, 2005; Logar and Pollock, 2005; Toldrá and Reig, 2006; Müller and Steinhart, 2007; see Chapters 22 and 26, this volume). There is increasing media coverage of food-related topics but, for a variety of reasons, media reporting may be incomplete or inaccurate (Butterworth, 2004; Logar and Pollock, 2005; Bruhn and Earl, 2006). This gives cause for concern, espe-cially as adverse messages or negative press related to food safety can have a major influence on consumers’ purchasing decisions; unfavourable messages weigh much more heavily in influencing consumer decisions than do positive or favourable ones.

Recent examples of food-related topics covered in a negative light in news-papers, magazines and other media include ‘mad cow disease’ (BSE), use of hormones in livestock production, pesticide and other xenobiotic residues in foods, use of food additives (preservatives and colorants), the planting of GM crops and the raising of GM animals for human consumption. Thus, markets for fish and animal food products are frequently challenged with stories relating to ‘health scares’. For example, although the nutritional value and potential health benefits of eating fish are regularly stated and increased consumption of fish is advised by nutritionists, many people appear to have a greater aware-ness of the safety risks that may be linked to fish consumption (Huss et al.,2004; Verbeke et al., 2005). The latter seems to be the result of a greater mass-media focus on pollution issues and contamination of fish than on com-municating information about dietary recommendations and the potential

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health benefits of increased consumption of fish. Thus, fish and farm animal producers need to identify clear, solidly documented ‘health benefit’ messages that can be used to promote the marketing of fish and animal products and offset the negative images created by ‘health scare’ propaganda.

In the mind of the general public, transgenic technology and GMOs seem to be both food (nutritional) and animal welfare issues, although the link between the two may be rather tenuous. Although there is currently no scientific evi-dence for any hazards unique to GMOs, there are consumer fears about food safety, ecosystem disturbance and degradation and animal welfare (Beardmore and Porter, 2003; Bennett et al., 2005):

● Food safety: is consumption of GMOs safe or are there unknown dangers?● Environmental impacts: how might escapees influence natural populations

of conspecifics and/or other species?● Welfare and ethics: is it morally defensible to manipulate higher organisms

to the extent possible using modern biotechnological techniques?

There are probably many reasons for the lack of clarity around the issue of GMOs, but one result of the confused state of affairs has been market resist-ance against such products (Tucker, 2003; Aerni, 2004; Kuiper et al., 2004; Deisingh and Badrie, 2005; Lehrer and Bannon, 2005; Logar and Pollock, 2005; Myhr and Dalmo, 2005; Tenbült et al., 2005; Bruhn and Earl, 2006; Dalal et al., 2006; Saher et al., 2006). In general, attitudes towards GMOs and organically grown food (OGF) are negatively related (Bennett et al., 2005; Dreezens et al., 2005; Saher et al., 2006). Strictly speaking, GMOs and OGFs refer to different methods of production rather than to any distinct properties or unique nutritional qualities of the end product. Nevertheless, consumers tend to treat such products as being inherently different from each other and from foods produced by conventional methods. The general public usually associates OGFs with ‘naturalness and pureness’, whereas GMOs are seen as having been altered extensively and manipulated by human hands, and this often results in a negative attitude towards GM products.

Consumer acceptance of a GM product also appears to be influenced by the way the generic product type is perceived; GM is less acceptable when a product type is viewed as being ‘natural and healthy’, such as the case with fruits and vegetables, than when the product is already seen as being ‘manufac-tured, unnatural and unhealthy’, as with cakes, biscuits and snack-foods (Tenbült et al., 2005). The resistance to GMOs appears to be particularly widespread with regard to farmed fish; this is not surprising, given that fish and fish prod-ucts would normally be considered as falling within the category of ‘natural and healthy’ foods. In other words, potential consumers often view GM farmed fish as being contrary to the image of an ‘ecologically sound, natural, health food’ (Bennett et al., 2005; Logar and Pollock, 2005).

Given this, there is a need for caution and increased transparency associ-ated with the regulation, licensing and broader introduction of such products, and it is also essential that effective methods be developed for the detection of GM products that enable them to be distinguished readily from non-GM entities (Beardmore and Porter, 2003; Kok and Kuiper, 2003; Kuiper et al., 2004;

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Achievements and Challenges 23

Liu-Stratton et al., 2004; Deisingh and Badrie, 2005; Kato et al., 2005; Lehrer and Bannon, 2005; Myhr and Dalmo, 2005; Bruhn and Earl, 2006; Roy and Sen, 2006; Stewart, 2006; Müller and Steinhart, 2007). For exam-ple, regulations need to include consideration of the safeties of the inserted DNA and markers introduced to help in the identification of GMOs. There must also be evaluation of the potential toxicity and allergenicity of the gene products resulting from the introduction of novel DNA, and the possibility of horizontal gene transfer also needs to be considered.

1.8 Concluding Remarks

There has been a rapid increase in aquaculture production over the course of the past few decades (http://www.fao.org/figis) and many predict that this expan-sion will continue for several years to come. If the predictions are to come to frui-tion, several prerequisites must be fulfilled. Government agencies and public authorities must create conditions that foster advances in aquaculture R & D. They should also promote advisory services designed to serve the industry and they must issue the regulations necessary for the maintenance of the trust and confidence of the public. Research institutions currently have a range of modern techniques available that enable them to provide results in genetics and selective breeding, nutrition and aqua-feed development and management practices that can lead to the production of quality aquaculture products at affordable cost. The industry must be in a position to realize the potential of the research results in a commercial setting and farmers must be receptive to, and apply, new techniques that lead to improvements in the efficiency of their operations.

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