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Fish Pathology E-Book

Ronald J. Roberts

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Beschreibung

Fish Pathology is the definitive, classic and essential book on the subject, providing in-depth coverage across all major aspects of fish pathology.

This new, fully updated and expanded fourth edition builds upon the success of the previous editions which have made Fish Pathology the best known and most respected book in the field, worldwide.Commencing with a chapter covering the aquatic environment, the book provides comprehensive details of the anatomy and physiology of teleosts, pathophysiology and sytematic physiology, immunology, neoplasia, virology, parasitology, bacteriology, mycology, nutritional pathology and other non-infectious diseases. A final chapter provides extremely useful details of the most widely-used and trusted laboratory methods in the area. Much new infomation is included in this new edition, including enhanced coverage of any diseases which have become commercially significant since publication of the previous edition

Beautifully illustrated in full colour throughout with many exceptional photographs, Fish Pathology, Fourth Edition, is an essential purchase for fish pathologists, fish veterinarians, biologists, microbiologists and immunologists, including all those working in diagnostic services worldwide. Personnel working in fish farming and fisheries will also find much of great use and interest within the book's covers. All libraries in universities and research establishments where biological and veterinary sciences are studied and taught should have copies of this landmark publication on their shelves.

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Veröffentlichungsjahr: 2012

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Table of Contents

Cover

Dedication

Title page

Copyright page

Contributors

Preface

1 The Aquatic Environment

INTRODUCTION

PHYSICAL AND CHEMICAL ASPECTS OF WATER QUALITY

ADVERSE BIOLOGICAL FACTORS FOR FISH HEALTH

CHARACTERISTICS OF DIFFERENT TYPES OF WATER

2 The Anatomy and Physiology of Teleosts

INTRODUCTION

THE INTEGUMENTARY SYSTEM

THE MUSCULOSKELETAL SYSTEM

THE RESPIRATORY SYSTEM

THE CIRCULATORY SYSTEM

BLOOD COMPOSITION

HAEMOPOIETIC TISSUE

THE RETICULOENDOTHELIAL SYSTEM

THE RENAL AND EXCRETORY SYSTEMS

THE DIGESTIVE SYSTEM

NUTRITION, METABOLISM AND GROWTH

THE REPRODUCTIVE SYSTEM

THE NERVOUS SYSTEM

SWIM-BLADDER

THE ENDOCRINE SYSTEM

3 The Pathophysiology and Systematic Pathology of Teleosts

PATHOPHYSIOLOGY

SYSTEMATIC PATHOLOGY

4 The Immunology of Teleosts

INTRODUCTION

NONSPECIFIC DEFENCE MECHANISMS

SPECIFIC DEFENCE MECHANISMS

5 Neoplasia of Teleosts

TUMOURS OF EPITHELIAL CELL ORIGIN

TUMOURS OF MESENCHYMAL ORIGIN

TUMOURS OF HAEMOPOIETIC ORIGIN

TUMOURS OF NERVE CELL ORIGIN

OTHER TUMOURS

6 The Virology of Teleosts

GENERAL BIOLOGY OF VIRUS INFECTIONS

PRACTICAL ASPECTS OF FISH VIROLOGY

VIRUS INFECTIONS OF FISH – DNA VIRUSES

RNA VIRUSES

7 The Parasitology of Teleosts

TAXONOMY OF FISH PARASITES

LIFE CYCLES OF PARASITES

PARASITES OF THE INTEGUMENT

PARASITES OF THE EYE

PARASITES OF THE VASCULAR SYSTEM

PARASITES OF THE CENTRAL NERVOUS SYSTEM

PARASITES OF THE SKELETAL SYSTEM

PARASITES OF THE VISCERA AND MUSCULATURE

PARASITES OF THE ALIMENTARY CANAL

ZOONOSES

8 The Bacteriology of Teleosts

THE FISH PATHOGENIC BACTERIA

9 The Mycology of Teleosts

OOMYCETES

CHYTRIDIOMYCETES (TRUE FUNGI)

ZYGOMYCETES (TRUE FUNGI)

DEUTEROMYCOTINA (Fungi imperfecti)

ALGAE

10 The Nutritional Pathology of Teleosts

ABSOLUTE NUTRITIONAL DEFICIENCY: STARVATION

DEFICIENCIES AND IMBALANCES OF MAJOR DIETARY COMPONENTS

TOXIC COMPONENTS OF THE DIET

TOXIC ORGANIC COMPOUNDS IN THE DIET

NUTRITIONAL CATARACTS

SINGLE-CELL PROTEIN TOXICITY

DRUG TOXICITY

11 Miscellaneous Non-infectious Diseases

GAS-BUBBLE DISEASE

LOW-TEMPERATURE DISEASES

WATER-BORNE IRRITANTS

BLUE-SAC, WHITE-SPOT AND YOLK SAC DEFORMITY DISEASE OF LARVAE

COLOURATION ANOMALIES

PHYSICAL DEFORMITIES

EARLY LIFE STAGE MORTALITY SYNDROME

CYSTIC CONDITIONS

TRAUMATIC INJURIES

JELLYFISH STING

SUNBURN

LIGHTNING STRIKE

ULCERATIVE DERMAL NECROSIS (UDN)

ACUTE ANAPHYLAXIS

12 Laboratory Methods

HEALTH CERTIFICATION

LABORATORY TESTING PROCEDURES

HISTOPATHOLOGY

HISTOPATHOLOGY: TECHNIQUES AND FORMULAE

TRANSMISSION AND SCANNING ELECTRON MICROSCOPY

BACTERIOLOGY

MYCOLOGY

PARASITOLOGY

PARASITOLOGY: TECHNIQUES AND FORMULAE

VIROLOGY

SEROLOGY

References

Fish Species Index

Subject Index

Dedication

To my long suffering wife Helen Macgregor. Her friendship, loyalty and support, when times were good and, more importantly, when they most decidedly were not, will always be appreciated.

This edition first published 2012 © 2012 by Blackwell Publishing Ltd.

Third edition © 2001 Harcourt Publishers Ltd.

Blackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’s publishing program has been merged with Wiley’s global Scientific, Technical and Medical business to form Wiley-Blackwell.

First published 1978

Second edition 1989

Third edition 2001

Fourth Edition 2012

Several editions also published in French, Spanish, Italian, German, Japanese, Mandarin and Bahasa Malay.

Registered office: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK

Editorial offices: 9600 Garsington Road, Oxford, OX4 2DQ, UK

The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK

2121 State Avenue, Ames, Iowa 50014-8300, USA

For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell.

The right of Ronald J. Roberts to be identified as the author of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988.

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, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher.

Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent professional should be sought.

Library of Congress Cataloging-in-Publication Data

Fish pathology / edited by Ronald J. Roberts. – 4th ed.

p. cm.

 Includes bibliographical references and index.

 ISBN 978-1-4443-3282-7 (hard cover : alk. paper)

 ISBN 978-1-1182-2293-5 (epdf)

 ISBN 978-1-1182-2296-6 (epub)

 ISBN 978-1-1182-2295-9 (mobi)

 I. Roberts, Ronald J., 1941–

 SH171.F58 2012

 597–dc23

2011035802

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

Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books.

Contributors

A.E. Ellis, MA (Oxon), PhD (Aberdeen)

Formerly Senior Principal Scientific Officer

Fisheries Research Service

Marine Laboratory

Aberdeen, Scotland

R.W. Hardy, BSc, PhD (Washington)

Director

Hagerman Center for Sustainable Aquaculture

University of Idaho

Idaho, United States

E.S. Munro, BSc (RGU, Aberdeen)

Group Leader Virology

Marine Scotland

Marine Laboratory

Aberdeen, Scotland

R.J. Roberts, BVMS, PhD (Glasgow)

FRCVS, FRCPath, FSB, FRSE, Commander of the Most Noble Order of the Crown (Thailand)

Hagerman Distinguished Visiting Professor

University of Idaho, United States

Technical Director

Landcatch Ltd

Ormsary, Argyll, Scotland

Emeritus Professor

formerly Founder Director

Institute of Aquaculture

University of Stirling, Scotland

H.D. Rodger, BVMS (Glasgow), MSc, PhD (Stirling)

MRCVS

Practice Principal

Vet-Aqua International

Oranmore, Co.

Galway, Ireland

C.J. Secombes, BSc (Leeds), PhD (Hull), DSc (Aberdeen), FSB, FRSE

Head, Scottish Fish Immunology Research Centre

University of Aberdeen

Scotland

D.A. Smail, BA (Oxon), MSc (Birmingham), PhD (CNAA)

Group Leader, Immunology & Infection

Marine Scotland

Marine Laboratory

Aberdeen, Scotland

R. Wootten, BSc, PhD (London)

Deputy Director

Institute of Aquaculture

University of Stirling

Stirling, Scotland

In addition, material from previous editions contributed by Dr A.M. Bullock, Dr G.N. Frerichs, Dr P. Tytler and Dr A.L.S. Munro has also been edited, updated and incorporated into the text.

Preface

This is the fourth edition of Fish Pathology. Little did I think when I penned the first few sentences in Nikola Fijan’s Laboratory in Zagreb, in 1976, that this book would survive to a fourth edition and be translated into nine languages. To my surprise and immense pleasure, it is now into its fourth decade and very much accepted as a standard work on its subject.

This edition has been revised and reset, incorporating much new information, and I am very grateful to Wiley-Blackwell for their encouragement to complete what has become a very large text and also for accepting my major demands for colour illustrations. The ability to use such illustrations throughout the text makes the task of writing very much easier, and this volume is illustrated to a degree which would have been unthinkable when planning that first edition, in early 1976.

Much of the revision and editing has once again been carried out in the beautiful and stimulating environment of the Hagerman Laboratories of the University of Idaho. Here, for the third time in my career, I have been presented with the opportunity to develop an international fish disease laboratory from the drawing board, as part of the new facility which Professor Hardy now directs. The facility provides an excellent resource for both cutting-edge research and more contemplative writing of texts such as this. Few can have been so fortunate, and I am most grateful to Professor Hardy and the University of Idaho for their generous support.

Teleost fish inhabit what is to most of us an alien world. Their exploitation for food or for sport has, until comparatively recently, been completely dependent on the hunting of wild stocks. Since the first edition of this book was published in 1978, however, the role of aquaculture in relation to world fish production has been transformed beyond measure. In 1978 aquaculture was responsible for less than 5% of the total fish consumed. Now the figure is nearer 50% by weight and more than 50% in terms of economic value. The blue revolution, no less significant than the green one, which led to self-sufficiency in rice and grain production in the developing world, is very much underway.

Control of fish diseases has been one of the major factors in allowing this growth in farmed fish production to take place. Our knowledge of fish pathology has expanded dramatically over the past decade, underpinned by the remarkable developments that have taken place in molecular biology and immunopathology. This has made production of a single text, which attempts to encapsulate the main aspects of all of the different disciplines of which the subject is comprised, rather challenging. This will, I suspect, be the last time that anyone will attempt such an all-embracing task. The individual subjects which make up fish pathology have now themselves grown to a size where they each demand their own monographs.

At least, however, I hope that this volume will, for a time, help to ensure that as aquaculture continues to grow, we will still be able to define, to diagnose and ultimately to control, in a sustainable way, the new diseases which will assuredly manifest themselves. The health and welfare of our fish stocks, both wild and farmed, are, I believe, measures of the quality of our entire aquatic environment.

The study of fish diseases requires a wide knowledge, not only of the potential pathogens but also of the environmental constraints and specialist adaptations which govern the ectothermic, aqueous existence of the teleosts. The inflammatory and immune responses of fishes are greatly modified by the nature of their environment. These, in turn, influence the epizootiology and the clinical characteristics of the various conditions and the methods by which they can be controlled.

One of the most heartening advances in the decade since the last edition of Fish Pathology was published has been the further development of sophisticated and efficient vaccines for many of the most damaging bacterial diseases. The advent of DNA and recombinant vaccines and immune enhancers has contributed greatly to fish welfare. Even more dramatic over the period has been the contribution of molecular biology to diagnostic advances such as those presented by real-time, or quantitative polymerase chain reaction (PCR), and the advent of breeding programmes that utilise molecular markers for genetic selection for disease resistance.

A very significant challenge presented in writing this book compared to the first three editions has been the advent of molecular techniques for the demonstration of relatedness between microorganisms. As a result of this, the taxonomy of many of the parasite and fungal groups has been completely rewritten. In the past, morphology and other phenotypic characteristics were dominant in deciding to which group a particular pathogen should be assigned. Now this can be decided with much greater certitude based on DNA homologies. While this has contributed greatly to precision, it has in many cases completely overturned previous assumptions. The taxonomies found therefore in the chapters on parasitology and mycology of fishes (Chapters 7 and 9, respectively) will appear somewhat strange to the reader accustomed to these previous certainties. I am very grateful to Dr Rod Wootten for his efforts to present a comprehensible taxonomic basis for the protistan parasites in particular and also to Dr Pieter Van West for his explanation to me of the new groupings of the Oomycetes and the true fungi.

Fish pathology is clearly a multidisciplinary field. The essential purpose of this book, like its predecessors, is to provide a corpus of basic information, drawn from these disciplines, which will be of value to the veterinarian, the microbiologist, the parasitologist, the nutritionist or the hydrologist. There are some 17 000 species of teleosts, all with specific adaptations at the gross, cellular and molecular levels, for their particular niche. In a book such as this, therefore, rather sweeping generalisations are sometimes necessary to render basic information more understandable.

I am very grateful to the authors of the contributed chapters for their forbearance, once again, and for their willingness to accept any editorial liberties that I may have taken in rewriting them for the sake of standardisation. I firmly believe that this is what gives the book its homogeneity and hope that they consider that their efforts have been justified. I must also acknowledge the assistance of Dr Emmett Shotts, formerly of the Leetown Laboratory, West Virginia and Dr Pieter van West and Professor Monty Priede of the University of Aberdeen for their willingness to review the bacteriology, mycology and anatomy chapters respectively, and Dr Hamish Rodger, formerly of the University of Pennsylvania, for general editorial support. I alone, however, am responsible for all errors and omissions.

Wiley-Blackwell, which also publishes the Journal of Fish Diseases, which I edit with Dr Wootten, has again generously allowed publication of many illustrations which first appeared in that journal. I am also grateful to the plethora of friends throughout the world who have made their illustrations available to me. Their generosity is acknowledged in the text wherever possible, but I apologise in advance for any omissions.

The preparation of this book would have been impossible without the drive, help and encouragement of Nigel Balmforth of Wiley-Blackwell, his assistant Carys Williams and the project manager Mirjana Misina. It was also heavily dependent on the encouragement and support of my colleagues Dr Laird Noh, Tracy Brown and Jana Cole at the University of Idaho and Sir William Lithgow, Hugh Currie, Alan Stewart and Neil Manchester at Landcatch Ltd. It is also important to acknowledge once again the determined efforts of Andrew Millar and K.G. Clarke, former colleagues at the University of Stirling, who so wholeheartedly relieved me of administrative duties so that I have, as Emeritus Professor, been able to concentrate on the real work of scientific research instead of the tyranny of the bean counter’s spread sheet.

Sadly during the course of the writing of this edition, Dr Tony Ellis, my first PhD student, who went on to establish an outstanding career in fish immunology and had contributed to every edition of this book, succumbed to cancer. He was a free spirit with a very original mind and an excellent scientist and teacher. He will be greatly missed.

Finally I have to thank my wife Helen for providing the line drawings and for her encouragement in what has certainly been a much more demanding task than either of us envisaged.

Ronald J. Roberts

Hagerman, Idaho

1

The Aquatic Environment

INTRODUCTION

Disease, in fishes, is closely linked to environmental stress. In the wild, they generally have some degree of freedom to modify their environment. They can move to more suitable conditions if faced with a negative environmental change such as a reduction in oxygen level or increase in temperature. Infected fish will even move to a warmer area to create an enhanced body temperature as an aid to increasing the rate of inflammatory response. In culture conditions, on the other hand, they have limited opportunity to choose their external environmental conditions. It is thus important that the conditions under which they are held provide environmental parameters suitable for all of the requirements of the particular species.

The aquatic environment encompasses a wide variety of features, virtually all of which influence the maintenance of homeostasis, essential for growth and reproduction of fishes. If altered beyond acceptable limits, they may predispose to, or actually cause, a wide range of disease processes. Among the most important environmental parameters are physical factors such as the temperature, the intensity and periodicity of light (including shading and background hue), the chemical composition of the water and its biological content, the availability of space and food and the frequency of fright stimuli such as moving shadows (EFSA 2008). Another important factor for wild fish and those farmed in extensive systems is the productivity of the ecosystem which sustains their food supply. Since consideration of ecosystems and the ecophysiological requirements of fish is beyond the scope of the present text, the reader is referred to Rankin and Jensen (1993), Macan (1974) and Odum (1971).

PHYSICAL AND CHEMICAL ASPECTS OF WATER QUALITY

TEMPERATURE

Fish have upper and lower thermal tolerance limits and optimum temperatures for growth, egg incubation, food conversion and resistance to specific diseases. These optima vary with species and may be different for different parameters such as oxygen tension and water pH. Fresh waters are subject to temperature fluctuations of up to 40°C caused by latitude, season, altitude, time of day, depth and other factors. The range of temperature change of sea-waters is much less, due to water circulation in the seas and oceans and the large volumes of water involved.

Many fish diseases are temperature modulated, with pathogenicity closely related to a specific temperature range. In many host–pathogen systems there is a balance between the host’s defences and the pathogen’s invasiveness, but this is readily modified by temperature change, especially if it is rapid.

Water temperature also affects other properties of the aquatic environment important for fish health. Dissolved gases generally decrease in solubility with increasing temperature (Table 1.1), whereas the solubility of toxic compounds, which are only sparingly soluble in water, such as crude oil and pesticides, increases with temperature rise. The toxicity of some substances such as heavy metals also increases with temperature (Wedemeyer 1997).

Table 1.1 Solubility of oxygen in water exposed to water-saturated air* (mg/litre).

*Values are quoted for 760 mmHg pressure. Under any other barometric pressure, P, the solubility S′ (mg/litre) is given by

where S is the solubility at 760 mmHg and p is the pressure (mmHg) of saturated water vapour at the prevailing temperature.

LIGHT

Light is a complex ecological factor whose components include colour spectrum (quality), intensity (quantity) and photoperiod (periodicity). The aquatic environment has peculiar and extremely variable characteristics, and ‘receptivity’ of fish to light changes profoundly from one species to another and, within the same species, from one developmental stage to another (Boeuf & Falcon 2001).

In natural waters and extensive farming systems, light levels can be changed only indirectly by methods such as increasing water depth and controlling unicellular algae, macrophytes and tree shade. Poor light penetration caused by absorbent or reflecting pollutants, such as clays, coal washings and paper wastes, diminishes algal productivity and may decrease the available levels of food for fish.

In intensive systems the light regime – intensity, photoperiod, shaded areas and light absorption by background – is more readily controlled. All of these parameters may contribute to aspects of the growth and maturation rate of fish. In such intensive culture systems, however, there is increasing evidence to suggest that ultraviolet light from excessive sunlight can result in sunburn of the dorsal surface, head or fins (Bullock 1987). With the development of thinning of the ozone layer as a result of excessive release of chlorinated compounds into the atmosphere, the problem of sunburn has become particularly serious in fish reared in the southern hemisphere. In addition, under certain conditions, even low levels of ultraviolet radiation can induce sunburn effects. This condition, known as photosensitisation, usually results from the incorporation in the diet of photoactive chemicals, often derived from drugs incorporated for therapeutic purposes or specific feed components such as porphyrins. In many cultured species, appropriate circadian periods of darkness and light are needed for proper timing and completion of events such as smoltification and sexual maturation, and good welfare and homeostasis often depend on appropriate photoperiod signals (Fjelldal, et al. 2005; Berril, et al. 2006).

PHYSICOCHEMICAL PARAMETERS

The Ionic Product of Water

Water behaves as a weak base and acid in that it is capable of losing or gaining a proton:

The equilibrium constant for the dissociation of water is given by

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