Mucosal Health in Aquaculture Page left intentionally blank Mucosal Health in Aquaculture Edited by Benjamin H. Beck Stuttgart National Aquaculture Research Center, Stuttgart, Arkansas, USA Eric Peatman School of Fisheries, Aquaculture, and Aquatic Sciences, Auburn University, Alabama, USA AMSTERDAM • BOSTON • HEIDELBERG • LONDON • NEW YORK OXFORD • PARIS • SAN DIEGO • SAN FRANCISCO • SINGAPORE SYDNEY • TOKYO Academic Press is an Imprint of Elsevier Academic Press is an imprint of Elsevier 125, London Wall, EC2Y 5AS, UK 525 B Street, Suite 1800, San Diego, CA 92101-4495, USA 225 Wyman Street, Waltham, MA 02451, USA The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, UK Copyright © 2015 Elsevier Inc. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means electronic, mechanical, photocopying, recording or otherwise without the prior written permission of the publisher. Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK: phone (+44) (0) 1865 843830; fax (+44) (0) 1865 853333; email: [email protected]. Alternatively, visit the Science and Technology Books website at www.elsevierdirect.com/rights for further information. Notice No responsibility is assumed by the publisher for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions or ideas contained in the material herein. Because of rapid advances in the medical sciences, in particular, independent verification of diagnoses and drug dosages should be made. British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication Data A catalog record for this book is available from the Library of Congress ISBN: 978-0-12-417186-2 For information on all Academic Press publications visit our website at http://store.elsevier.com/ Typeset by Thomson Digital Printed and bound in the United States of America Dedication We dedicate this text to fish that have captivated us since youth. Slimy and colorful. Resilient and enduring. Fried, broiled, and blackened. Page left intentionally blank Contents List of contributors xi Acknowledgments xiii 1 Why mucosal health? 1 Eric Peatman; Benjamin H. Beck 1.1 Why mucosal health? 1 2 Overview of fish immunity 3 Rosario Castro; Carolina Tafalla 2.1 Introduction 4 2.2 Organs, tissues, and general structures 6 2.3 Cellular components 10 2.4 Soluble mediators of immunity 21 2.5 Immune mechanisms 30 Acknowledgments 36 References 36 3 Overview of mucosal structure and function in teleost fishes 55 Tracy S. Peterson 3.1 Introduction 55 3.2 Integument 58 3.3 Intestine 60 References 63 4 Fish mucosal immunity: skin 67 María Ángeles Esteban; Rebeca Cerezuela 4.1 Introduction 67 4.2 The mucous layer as a barrier against pathogens 69 4.3 Resident cell types found in the skin 72 4.4 Skin immune responses in diseased fish 74 4.5 Future directions and conclusions 87 References 88 5 Fish mucosal immunity: gill 93 Erling Olaf Koppang; Agnar Kvellestad; Uwe Fischer 5.1 Introduction 94 5.2 Embryology and anatomy 95 viii Contents 5.3 Vascular system 97 5.4 Epithelium 98 5.5 Blood cells and other immune cells 101 5.6 Mucosal immunity and the gills 110 5.7 Infectious salmon anemia 113 5.8 Red mouth disease (yersiniosis) 115 5.9 Amoebic gill disease 116 5.10 Ichthyophthiriasis (“white spot disease”) 117 5.11 Concluding remarks 120 References 120 6 Fish mucosal immunity: intestine 135 Irene Salinas; David Parra 6.1 Introduction 136 6.2 The mucus layer as a barrier against mucosal pathogens 137 6.3 Resident cell types found in the gastrointestinal tract of teleosts 140 6.4 Gut immune responses in diseased fish 145 6.5 The embryology and ontogeny of the gut immune system 152 6.6 Laboratory models that can contribute to our knowledge of gut immunity in aquaculture fish 155 6.7 Concluding remarks 158 References 158 7 Environmental impacts on fish mucosa 171 Henrik Sundh; Kristina S. Sundell 7.1 Introduction 171 7.2 Intestinal barrier function 172 7.3 Intestinal barrier function and health 177 7.4 Environmental factors affecting the intestinal barrier 177 7.5 Host–pathogen interactions and intestinal barrier function 180 7.6 The effect of environmental salinity on the intestinal barrier 183 7.7 Mediators of a decreased barrier function 184 7.8 Concluding remarks 187 References 188 8 Nutritional impacts on fish mucosa: dietary considerations 199 Jesse Trushenski 8.1 Introduction 199 8.2 Lipids and gut health 200 8.3 Antinutritional factors and gut health 202 8.4 Concluding remarks 206 References 206 Contents ix 9 Nutritional impacts on fish mucosa: immunostimulants, pre- and probiotics 211 Christopher Marlowe A. Caipang; Carlo C. Lazado 9.1 Background 212 9.2 Immunostimulants 214 9.3 Prebiotics 221 9.4 Probiotics 229 9.5 Dietary administration of immunostimulants and their effects on mucosal immunity and disease resistance 241 9.6 Dietary administration of prebiotics and their effects on mucosal immunity 243 9.7 Dietary administration of probiotics and their effects on mucosal immunity 249 9.8 Concluding remarks 254 Acknowledgments 255 References 255 10 The fish microbiome and its interactions with mucosal tissues 273 Daniel L. Merrifield; Ana Rodiles 10.1 Introduction 273 10.2 Ontology of the fish microbiome 274 10.3 Microbiota associated with the skin 276 10.4 Microbiota associated with the gills 280 10.5 The gut microbiomes of fish 281 10.6 Microbiome–mucosa interactions 287 10.7 Future research 289 References 289 11 Mucosal vaccines 297 Esteban Soto; Matt J. Griffin; Jaime A. Tobar 11.1 Introduction 297 11.2 Types of mucosal vaccines in aquaculture 300 11.3 Delivery of mucosal vaccines in aquaculture 309 References 318 12 Mucosal immunity in mollusks 325 Bassem Allam; Emmanuelle Pales Espinosa 12.1 Introduction 325 12.2 Molluscan mucosal epithelia 327 12.3 Mucus matrix composition and structure 330 12.4 The diverse functions of mollusk mucus 331 12.5 Host–microbe interactions at mucosal interfaces in metazoans 332 12.6 Mucosal immunity in mollusks 336 12.7 Immune recognition 337 12.8 Immune activation and cell signaling 342 x Contents 12.9 Effector molecules 342 12.10 Endocytic activity by epithelial cells 346 12.11 Hemocytes 347 12.12 Melanization and biomineralization 350 12.13 Enhancement of mucosal immunity 351 12.14 The promise of new technologies 352 12.15 Concluding remarks 352 Acknowledgments 353 References 353 13 New frontiers in mucosal health in aquaculture 371 Benjamin H. Beck; Eric Peatman 13.1 Next-generation sequencing-based approaches 372 13.2 Gene-editing 375 13.3 Conclusions 376 References 376 Subject Index 379 List of contributors Bassem Allam School of Marine and Atmospheric Sciences, Stony Brook University, Stony Brook, New York, USA María Ángeles Esteban Fish Innate Immune System Group, Department of Cell Biology and Histology, Faculty of Biology, Campus of International Excellence, Campus Mare Nostrum, University of Murcia, Murcia, Spain Benjamin H. Beck Stuttgart National Aquaculture Research Center, Stuttgart, Arkansas, USA Christopher Marlowe A. Caipang School of Applied Science, Temasek Polytechnic, Singapore Rosario Castro Centro de Investigación en Sanidad Animal (CISA-INIA), Valdeolmos, Madrid, Spain Rebeca Cerezuela Fish Innate Immune System Group, Department of Cell Biology and Histology, Faculty of Biology, Campus of International Excellence, Campus Mare Nostrum, University of Murcia, Murcia, Spain Emmanuelle Pales Espinosa School of Marine and Atmospheric Sciences, Stony Brook University, Stony Brook, New York, USA Uwe Fischer Federal Research Institute for Animal Health, Friedrich-Loeffler Institut, Greifswald-Insel Riems, Germany Matt J. Griffin Thad Cochran National Warmwater Aquaculture Center, Aquatic Research and Diagnostic Laboratory, College of Veterinary Medicine, Mississippi State University, Stoneville, Mississippi, USA Erling Olaf Koppang Section of Anatomy and Pathology, Faculty of Veterinary Medicine and Biosciences, Norwegian University of Life Sciences, Oslo, Norway Agnar Kvellestad Section of Anatomy and Pathology, Faculty of Veterinary Medicine and Biosciences, Norwegian University of Life Sciences, Oslo, Norway Carlo C. Lazado Aquaculture Genomics Research Unit, Faculty of Biosciences and Aquaculture, University of Nordland, Bodø, Norway Daniel L. Merrifield School of Biological Sciences, Plymouth University, Plymouth, United Kingdom xii List of contributors David Parra Department of Cell Biology, Physiology and Immunology. Universitat Autonoma de Barcelona, Cerdanyola del Valles, Spain Eric Peatman School of Fisheries, Aquaculture, and Aquatic Sciences, Auburn University, Alabama, USA Tracy S. Peterson Department of Regulatory Science, University of Arkansas at Pine Bluff, Pine Bluff, Arkansas, USA Ana Rodiles School of Biological Sciences, Plymouth University, Plymouth, United Kingdom Irene Salinas Center for Evolutionary and Theoretical Immunology, Department of Biology, University of New Mexico, Albuquerque, New Mexico, USA Esteban Soto Department of Biomedical Sciences, Ross University School of Veterinary Medicine, West Farms, St. Kitts, West Indies Kristina S. Sundell Fish Endocrinology Laboratory, Department of Biological and Environmental Sciences, University of Gothenburg, Gothenburg, Sweden Henrik Sundh Fish Endocrinology Laboratory, Department of