Bacterial Fish Pathogens Brian Austin • Dawn A. Austin Bacterial Fish Pathogens Disease of Farmed and Wild Fish Sixth Edition Brian Austin Dawn A. Austin Institute of Aquaculture School of Life Sciences University of Stirling Heriot-Watt University Stirling , UK Edinburgh , Scotland , UK ISBN 978-3-319-32673-3 ISBN 978-3-319-32674-0 (eBook) DOI 10.1007/978-3-319-32674-0 Library of Congress Control Number: 2016952087 4th edition: © Praxis Publishing Ltd, Chichester, UK, 2007 5th edition: © Springer Science+Business Media Dordrecht 2012 © Springer International Publishing Switzerland 2016 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifi cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfi lms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specifi c statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. Printed on acid-free paper This Springer imprint is published by Springer Nature The registered company is Springer International Publishing AG Switzerland To Aurelia Jean Pref ace This sixth edition of Bacterial Fish Pathogens is the successor to the original ver- sion, fi rst published by Ellis Horwood Limited in 1987, which was planned to fi ll the need for an up-to-date comprehensive text on the biological aspects of the bacte- rial taxa which cause disease in fi sh. In those days, interest focused largely on bacte- rial kidney disease, the Cytophaga -like bacteria (CLBs), furunculosis and vibriosis (caused by Vibrio anguillarum). Over the intervening years, we have witnessed changes in impetus as new methodologies, notably involving molecular biology, have been developed. Interest in vaccines has been overtaken by probiotics and plant products. In terms of the number of publications, Aeromonas hydrophila is the pathogen of the moment. The impetus to prepare a sixth edition stemmed initially from discussion with the publisher when it became apparent that the book was par- ticularly well used (there have been >19,000 downloads since the publication of the fi fth edition in 2012). Since publishing the fi fth edition in 2012, there has been only a comparatively few new bacterial fi sh pathogens described, but numerous publica- tions deal with diagnosis, pathogenicity and especially control. Consequently with all the new publications, we considered that it is timely to prepare a new edition. Thinking back to the preparation of the fi rst edition, it is mind numbing how the mechanics of publication have changed. For the fi rst edition, we spent ages in librar- ies, using interlibrary loans; pouring over paper copies of journals, books and monographs; making handwritten notes; and preparing text for typing [on a type- writer – corrections led to use of white-out or the typing of replacement pages]. Line drawings were done using a drawing board, Rotring pens, set squares/French curves and stencils. Photographs were developed in a dark room and could be hit or miss in terms of quality. Now, we have access to an even wider range of journals electroni- cally, which have been accessed from the laptop computer via a wireless Internet connection. Weeks of waiting for interlibrary loans from around the globe did not even feature during the initial research phase of the fi fth or sixth editions. Because of the extensive new information published since the start of the millennium, we have been selective and, in particular, have once again condensed details of the pathology of the diseases, because there are excellent texts already available, which cover detailed aspects of the pathological conditions. For some diseases that have vii viii Preface occurred only occasionally and mostly in the distant past, we have condensed the text. Nevertheless, this sixth edition will hopefully meet the needs of the readership. As with all the preceding editions, it is emphasised that most of the information still appertains to diseases of farmed, rather than wild, fi sh. The scope of the book covers all of the bacterial taxa, which have at one time or another been reported as fi sh pathogens. Of course, it is realised that some taxa are merely secondary invaders of already damaged tissues, whereas others comprise serious, primary pathogens. Shortcomings in the literature or gaps in the overall understanding of the subject have been highlighted, sometimes with subtlety. As a general comment, it is still apparent that little is known about anaerobic or micro- aerophilic pathogens – does this mean that they do not exist, or are scientists ignor- ing them? Also, it is mostly unclear whether organisms act by themselves or with others to cause a disease manifestation. Uncultured pathogens have been recognised in the form of Candidatus whereby the organisms may be visualised in pathological material, but not grown in the laboratory. The cause of some diseases is uncertain, although in the case of red mark syndrome/strawberry disease of rainbow trout, there is serological and molecular evidence for the involvement of rickettsial-like organisms, but which are still yet to be cultured. One wonders how many times the presence of uncultured or diffi cult-to-culture organisms is masked by culturable contaminants or secondary invaders. A fi nal comment is that the overwhelming bulk of publications deal with diseases caused by single species of pathogen. Is this really true in that disease is a pure culture phenomenon, or is science missing micro- bial consortia that could work synergistically, simultaneously or sequentially to cause disease? In preparing the text, we have sought both advice and material from colleagues. We are especially grateful to the following for the supply of photographs: Dr. J.W. Brunt Dr. G. Dear Dr. M. Crumlish Professor H. Daskolov Dr. T. Itano Professor V. Jencic Dr. D.-H. Kim Professor C. Mothersill Dr. A. Newaj-Fyzul Dr. P. Orozova Dr. N. Pieters Dr. S.M. Sharifuzzaman Professor J.F. Turnbull Professor X.-H. Zhang Stirling, UK Brian Austin Scotland, UK Dawn A. Austin 2016 Contents 1 Introduction . 1 Conclusions . 19 References . 19 2 Gram-Positive Bacteria (Anaerobes and ‘Lactic Acid’ Bacteria) . 21 Anaerobes . 21 Clostridiaceae Representative . 22 Clostridium botulinum . 22 Eubacteriaceae Representative . 25 Eubacterium tarantellae . 25 Lactic Acid Bacteria . 27 Carnobacteriaceae Representative . 27 Carnobacterium piscicola (and the Lactobacilli) . 28 Leuconostocacceae Representative . 31 Weissella sp. 31 Weissella ceti . 33 Gram-Positive Cocci in Chains: The Early Literature . 34 General Comments . 34 Isolation . 36 Epizootiology . 36 Development of Vaccines . 37 Use of Inhibitory Compounds . 38 Enterococcaceae Representatives . 38 Enterococcus (Streptococcus) faecalis subsp. liquefaciens . 38 Vagococcus salmoninarum . 39 Streptococcaceae Representatives . 41 Lactococcus garvieae (= Enterococcus seriolicida) . 41 Lactococcus lactis subsp. lactis . 46 Lactococcus piscium . 47 Streptococcus agalactiae (= Str. diffi cilis) . 48 Streptococcus dysgalactiae subsp. dysgalactiae . 53 Streptococcus ictaluri . 55 ix x Contents Streptococcus iniae . 56 Streptococcus milleri . 63 Streptococcus parauberis . 64 Streptococcus phocae . 66 References . 68 3 Aerobic Gram-Positive Rods and Cocci . 83 Erysipelothrix rhusiopathiae . 83 Characteristics of the Disease . 83 Isolation . 83 Characteristics of the Pathogen . 84 Pathogenicity . 84 Disease Control . 84 Renibacterium salmoninarum . 84 Characteristics of the Disease . 84 Isolation . 86 Characteristics of the Pathogen . 89 Diagnosis . 93 Epizootiology . 98 Pathogenicity . 101 Disease Control . 103 Aerococcaceae Representative . 106 Bacillaceae Representatives . 107 Corynebacteriaceae Representatives . 110 Coryneform Bacteria . 112 Microbacteriaceae Representative . 113 Micrococcaceae Representative . 113 Mycobacteriaceae Representatives . 115 Nocardiaceae Representatives . 125 Planococcaceae Representative . 136 Staphylococcaceae Representatives . 138 References . 141 4 Aeromonadaceae Representatives (Motile Aeromonads) . 161 Aeromonas allosaccharophila . 162 Characteristics of the Disease . 162 Isolation . ..