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Fish Viruses and Bacteria: Pathobiology and Protection

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Taking a disease-based approach,æFish Viruses and Bacteria: Pathobiology and Protectionæfocuses on the pathobiology of and protective strategies against the most common, major microbial pathogens of economically important marine and freshwater fish.æThe book covers well-studied, notifiable piscine viruses and bacteria, including new and emerging diseases which can become huge threats to local fish populations in new geographical regions if transported there via infected fish or eggs. An invaluable bench book for fish health consultants, veterinarians and all those wanting instant access to information, this book is also a useful textbook for students specializing in fish health and research scientists initiating fish disease research programmes.

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1 Infectious Pancreatic Necrosis Virus



Infectious Pancreatic Necrosis Virus

Arun K. Dhar,1,2* Scott LaPatra,3 Andrew Orry4­ and F.C. Thomas Allnutt1


BrioBiotech LLC, Glenelg, Maryland, USA; 2Aquaculture Pathology Laboratory,

School of Animal and Comparative Biomedical Sciences, The University of

Arizona, Tucson, Arizona, USA; 3Clear Springs Foods, Buhl, Idaho, USA;


Molsoft, San Diego, California, USA

1.1  Introduction

Infectious pancreatic necrosis virus (IPNV), the aetiological agent of infectious pancreatic necrosis (IPN), is a double-stranded RNA (dsRNA) virus in the family Birnaviridae (Leong et  al., 2000; ICTV, 2014).

The four genera in this family include Aquabirnavirus,

Avibirnavirus, Blosnavirus and Entomobirnavirus

(Delmas et al., 2005), and they infect vertebrates and invertebrates. Aquabirnavirus infects aquatic species

(fish, molluscs and crustaceans) and has three species: IPNV, Yellowtail ascites virus and Tellina virus.


2 Infectious Haematopoietic Necrosis Virus



Infectious Haematopoietic

Necrosis Virus

Jo-Ann C. Leong1* and Gael Kurath2


Hawai‘i Institute of Marine Biology, University of Hawai‘i at Maˉnoa,

–ne‘ohe, Hawai‘i, USA; 2Western Fisheries Research Center, US


Geological Survey, Seattle, Washington, USA

2.1  Introduction

Infectious haematopoietic necrosis virus (IHNV, infectious hematopoietic necrosis virus) is a

Rhabdovirus that causes significant disease in Pacific salmon (Oncorhynchus spp.), Atlantic salmon

(Salmo salar), and rainbow and steelhead trout

(O. mykiss). The disease that it causes, infectious haematopoietic necrosis (IHN), was first detected in cultured sockeye salmon (O. nerka) in the Pacific

Northwest of North America and IHNV was first cultured in 1969 (see Bootland and Leong, 1999).

IHNV is the type species and reference virus for the

Novirhabdovirus genus of the family Rhabdoviridae.

The viral genome is a linear, single-stranded RNA


3 Viral Haemorrhagic Septicaemia Virus



Viral Haemorrhagic Septicaemia


John S. Lumsden*

Department of Pathobiology, Ontario Veterinary College, University of Guelph,

Guelph, Ontario, Canada and Adjunct Professor, Department of Pathobiology,

St. George’s University, True Blue, Grenada

3.1  Introduction

Viral haemorrhagic septicaemia virus (VHSV), the aetiological agent of viral haemorrhagic septicaemia (VHS) is a member of the family Rhabdoviridae within the order Mononegavirales. These viruses have single-stranded, negative-sense (nega) RNA genomes with enveloped bullet- or cone-shaped nucleocapsids. The economically most important pathogenic rhabdoviruses are VHSV, infectious haematopoietic necrosis virus (IHNV; both in the genus Novirhabdovirus) and spring viraemia of carp virus (SVCV; new name Carp sprivivirus, genus Sprivivirus). Please refer to the International

Committee on Taxonomy of Viruses for details

(ICTV, 2015). VHSV and IHNV share the same six genes that read from the 3′ end to the 5′ end of the genome as N (nucleocapsid protein), P (phosphoprotein), M (matrix protein), G (glycoprotein), NV


4 Epizootic Haematopoietic Necrosis and European Catfish Virus



Epizootic Haematopoietic Necrosis and European Catfish Virus

Paul Hick1, Ellen Ariel2 and Richard Whittington1*


Faculty of Veterinary Science, University of Sydney, Camden, New South

Wales, Australia; 2College of Public Health, Medical and Veterinary Sciences,

James Cook University, Townsville, Queensland, Australia

4.1  Introduction

Epizootic haematopoietic necrosis (EHN) disease is restricted to Australia and is caused by the ranavirus Epizootic haematopoietic necrosis virus

(EHNV). This systemic disease causes high mortality among naturally infected wild redfin perch

(Perca fluviatilis), has an impact on farmed rainbow trout (Oncorhynchus mykiss) and can threaten populations of native Australian fishes (Whittington et al., 2010; Becker et al., 2013). European catfish virus (ECV) is a closely related ranavirus that was originally isolated from sheatfish (Silurus glanis) in

Germany (Ahne et al., 1989). This virus causes high mortalities in sheatfish and black bullhead catfish


5 Oncogenic Viruses: Oncorhynchus masou Virus and Cyprinid Herpesvirus



Oncogenic Viruses: Oncorhynchus masou Virus and Cyprinid


Mamoru Yoshimizu,1* Hisae Kasai,1 Yoshihiro Sakoda,2

Natsumi Sano3 and Motohiko Sano4


Faculty of Fisheries Sciences, Hokkaido University, Minato, Hakodate, ­Japan;

Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, J­ apan;


Graduate School of Bioresources, Mie University, Tsu, Japan; 4Faculty of ­Marine

Science, Tokyo University of Marine Science and Technology, Tokyo, Japan


5.1  Introduction

Due to their distinctive appearance and obvious pathological nature, tumours of fish have been recognized for centuries. Publications on fish tumours are widely scattered in the scientific literature (Walker, 1969; Anders and Yoshimizu, 1994).

The largest registry of tumours in lower animals was established at the Smithsonian Institution in

Washington, DC, in 1965. Pathologies range from benign epidermal papillomas to metastatic melanomas and hepatocellular carcinomas in more than


6 Infectious Salmon Anaemia



Infectious Salmon Anaemia

Knut Falk* and Maria Aamelfot

The Norwegian Veterinary Institute, Oslo, Norway

6.1  Introduction

Infectious salmon anaemia (ISA) is a significant infectious viral disease of farmed Atlantic salmon,

Salmo salar L., that was first reported in Norway during 1984 (Thorud and Djupvik, 1988).

Outbreaks of ISA have an impact on the economy of the Atlantic salmon aquaculture industry, and this has led to the implementation of large-scale biosecurity measures. Outbreaks have now been reported in most Atlantic salmon farming areas, including the east coast of Canada and the USA,

Scotland, Norway, the Faroe Islands and Chile

(Rimstad et al., 2011). In Chile and the Faroe

Islands, the disease caused major economic setbacks and left the entire industry with an uncertain future (Mardones et al., 2009; Christiansen et al.,

2011) in a manner similar to that in Norway in and after 1989 (Håstein et al., 1999; Rimstad et al.,


7 Spring Viraemia of Carp



Spring Viraemia of Carp

Peter Dixon and David Stone*

Centre for Environment, Fisheries and Aquaculture Science (Cefas) Weymouth

Laboratory, Weymouth, UK

7.1  Introduction

Spring viraemia of carp (SVC) is often a fatal haemorrhagic disease of common carp, Cyprinus carpio, and other fishes. It is caused by the spring viraemia of carp virus (SVCV), a rhabdovirus (Fig. 7.1) of the type species, Carp sprivivirus, in the genus Sprivivirus

(Stone et  al., 2013; Adams et  al., 2014). SVC is reportable to the World Organisation for Animal

Health (OIE), but the virus is serologically related to other non-reportable viruses.

7.2  Hosts

SVC has been described in numerous hosts, although some early reports of the disease may actually have been carp erythrodermatitis, a bacterial disease. In addition to the common and koi carp

(C. carpio), hosts include the following cyprinids

(where no citation is referenced, check Dixon, 2008): bighead carp, Aristichthys nobilis; crucian carp,


8 Channel Catfish Viral Disease



Channel Catfish Viral Disease

Larry A. Hanson1* and Lester H. Khoo2


Department of Basic Sciences, College of Veterinary Medicine, Mississippi

State University, Mississippi,USA; 2Thad Cochran Warmwater Aquaculture

Center, Stoneville, Mississippi, USA

8.1  Introduction

Channel catfish viral disease (CCVD) is an acute viraemia that occurs primarily among young (0–4 month old) channel catfish (Ictalurus punctatus) in aquaculture. CCVD outbreaks occur almost exclusively in the summer when water temperatures exceed 25°C and may exceed 90% mortality in less than 2 weeks. Older fish may experience a more chronic outbreak, often with secondary Flavo­ bacterium columnare or Aeromonas infections that can mask the underlying CCVD (Plumb, 1978).

Pond-to-pond spread is often reported within fingerling production facilities. The disease was first described by Fijan et al. (1970) and the most notable clinical signs were exophthalmia, abdominal distension, disoriented swimming and rapidly increasing mortality.


9 Largemouth Bass Viral Disease



Largemouth Bass Viral Disease

Rodman G. Getchell1* and Geoffrey H. Groocock2


Veterinary Medical Center, Department of Microbiology and Immunology, Cornell ­University, Ithaca, New York, USA; 2Transit Animal Hospital, Depew,

New York, USA

9.1  Introduction

The first Ranavirus was isolated from leopard frogs, Rana pipiens, in the eastern USA (Granoff et al., 1965). Thirty years later, the first outbreak of largemouth bass viral disease was reported at

Santee-Cooper Reservoir, South Carolina by Plumb et al. (1996), who isolated the virus concerned on fathead minnow, Pimephales promelas (FHM) cells inoculated with filtered homogenates from two infected adult largemouth bass, Micropterus salmoides, collected during the mortality event.

Icosahedral virus particles (enveloped virions about

174 nm in diameter) were transmitted to uninfected largemouth bass by experimental transmission by injection. The isolate was tentatively classified as belonging to the family Iridoviridae and largemouth bass virus (LMBV) proposed as its name (Plumb et al., 1996).


10 Koi Herpesvirus Disease



Koi Herpesvirus Disease

Keith Way* and Peter Dixon

Centre for Environment, Fisheries and Aquaculture Science (Cefas)

Weymouth Laboratory, Weymouth, UK

10.1  Introduction

Koi herpesvirus disease (KHVD) is a herpesvirus infection (Hedrick et al., 2000) that induces a lethal acute viraemia that is highly contagious in common carp (Cyprinus carpio) and varieties of C. carpio such as koi carp and ghost carp (koi × common carp) (Haenen et al., 2004). The causative agent is classified as Cyprinid herpesvirus 3 (CyHV-3), a member of the family Alloherpesviridae and one of ten alloherpesviruses that infect fishes (Boutier et al., 2015a).

The transmission of CyHV-3 is horizontal and can occur directly or indirectly. Uchii et al. (2014) suggested that CyHV-3 in recovered fish reactivates periodically when the water temperature increases and transmits to naive fish when they are in close contact, such as at spawning. Direct transmission occurs by skin-to-skin contact between infected and naive carp, and through the cannibalistic and necrophagous behaviour of carp. Several vectors may facilitate the indirect transmission of CyHV-3.


11 Viral Encephalopathy and Retinopathy



Viral Encephalopathy and Retinopathy

Anna Toffan*

OIE Reference Centre for Viral Encephalopathy and Retinopathy,

Istituto Zooprofilattico Sperimentale delle Venezie, Legnaro (Padova), Italy

11.1  Introduction

Viral encephalopathy and retinopathy (VER), also known as viral nervous necrosis (VNN) is a severe neuropathological disease caused by RNA viruses of the genus Betanodavirus (Family: Nodaviridae).

This infectious agent, detected in the late 1980s, spread worldwide, became endemic and came to represent a major limiting factor for mariculture in several countries. The disease has recently been included among the most significant viral pathogens of finfish, given the expanding host range and the lack of properly effective prophylactic measures

(Rigos and Katharios, 2009; Walker and Winton,

2010; Shetty et al., 2012).

11.2  The Infectious Agents

The causative agent of the disease is a small

(25–30 nm diameter), spherical, non-enveloped virion, with a bi-segmented genome made of two singlestranded positive-sense RNA molecules. The name


12 Iridoviral Diseases: Red Sea Bream Iridovirus and White Sturgeon Iridovirus



I�ridoviral Diseases: Red Sea

Bream Iridovirus and White

Sturgeon Iridovirus

Yasuhiko Kawato,1 Kuttichantran Subramaniam,2

Kazuhiro Nakajima,1 Thomas Waltzek2 and Richard



National Research I­nstitute of Aquaculture, Japan Fisheries Research and

Education Agency, ­Nakatsuhamaura, Minami-Ise, Mie, Japan; 2Department of

Infectious D

­ iseases and Pathology, University of Florida, Gainesville, Florida,

USA; 3Faculty of Veterinary Science, University of Sydney, Camden, New South

Wales, Australia

12.1  Red Sea Bream Iridovirus

12.1.1  Introduction

The red sea bream iridovirus (RSIV) has a doublestranded DNA genome in an icosahedral virion capsid that is 200 nm in diameter. According to the

International Committee on the Taxonomy of

Viruses (ICTV), it is in the genus Megalocytivirus within the family Iridoviridae, although it has not yet been approved as a species within that genus


13 Alphaviruses in Salmonids



Alphaviruses in Salmonids

Marius Karlsen1* and Renate Johansen2


PHARMAQ AS, Oslo, Norway; 2PHARMAQ Analytiq, Bergen, Norway

13.1  Introduction

Alphavirus is a genus of RNA viruses belonging to the family Togaviridae. Most known alphaviruses are mosquito borne and cause diseases in terrestrial hosts such as birds, rodents and larger mammals, including humans (Strauss and Strauss, 1994).

Infections may lead to diverse symptoms, such as rashes, gastrointestinal problems, arthritis/muscular inflammation and encephalitis (Kuhn, 2007; Steele and Twenhafel, 2010). Salmon pancreas disease virus, which is commonly named Salmonid alphavirus and abbreviated SAV (Weston et  al., 2002), is the only known alphavirus that has fish as a natural host (Powers et al., 2001). SAV is distantly related to other members of the genus, but it still causes pathology that may resemble some of that seen in mammals (McLoughlin and Graham, 2007; Biacchesi et al., 2016). The first isolation of SAV in cell culture was reported in 1995 from marine farmed


14 Aeromonas salmonicida and A. hydrophila



Aeromonas salmonicida and A. hydrophila

Bjarnheidur K. Gudmundsdottir1* and Bryndis



Faculty of Medicine, University of Iceland, Reykjavik, Iceland;

Matís, Reykjavik, Iceland


14.1  Introduction

Aeromonas belongs to the family Aeromonadales within the class Gammaproteobacteria (Colwell et al.,

1986). Aeromonads occur in freshwater, estuarine and marine environments, invertebrates, vertebrates and soils (Janda and Abbott, 2010). The type species is the motile A. hydrophila, an animal pathogen; in contrast, the species A. salmonicida, a fish pathogen, is non-motile. Aeromonads induce furunculosis, atypical furunculosis, ulcerative diseases, motile

Aeromonas septicaemia (MAS) and tail and fin rot in fishes (Cipriano and Austin, 2011). A. hydrophila and other motile species (e.g. A. veronii biovar. sobria, A. bestiarum, A. dhakensis) cause diseases in aquaculture and are potentially zoonotic pathogens (Rahman et  al., 2002; Janda and Abbott,


15 Edwardsiella spp.



Edwardsiella spp.

Matt J. Griffin,* Terrence E. Greenway and David J. Wise

Thad Cochran National Warmwater Aquaculture Center,

Mississippi State University, Stoneville, Mississippi, USA

15.1  Introduction

The Edwardsiella (family Enterobacteriacae) was originally described as a new genus of the Enter­ obacteriaceae in the mid-1960s; it represented 37 isolates recovered from open wounds, blood, urine and faeces of humans and animals in the USA,

Brazil, Ecuador, Israel and Japan (Ewing et  al.,

1965). In spite of this, the species of Edwardsiella are mostly considered to be pathogens of fish

(Mohanty and Sahoo, 2007; Table 15.1). E. tarda was first reported from outbreaks in farmed channel catfish (Ictalurus punctatus) in Arkansas in the

USA (Meyer and Bullock, 1973) and has become one of the most globally recognized fish pathogens, affecting both wild and cultured fish worldwide

(Park et al., 2012).

Similarly, E. ictaluri was described from farmraised catfish in the south-eastern USA in the early


16 Flavobacterium spp.



�Flavobacterium spp.

Thomas P. Loch* and Mohamed Faisal

Department of Pathobiology and Diagnostic Investigation, College of Veterinary

Medicine, Michigan State University, East Lansing, Michigan, USA

16.1  Introduction

Flavobacterial diseases in fish are mainly attributed to three Gram-negative, yellow-pigmented bacteria: Flavobacterium psychrophilum, the cause of bacterial cold water disease (BCWD) and rainbow trout fry syndrome (RTFS; Davis, 1946; Borg,

1948; Holt, 1987; Bernardet and Grimont, 1989);

F. columnare, the cause of Columnaris disease (CD;

Davis, 1922; Ordal and Rucker, 1944; Bernardet and Grimont, 1989); and F. branchiophilum, the putative agent of bacterial gill disease (BGD;

Wakabayashi et al., 1989).

16.1.1  Flavobacterium psychrophilum

Description of the microorganism

Borg (1948) described epizootics at water temperatures of 6–10°C in farmed coho salmon

(Oncorhynchus kisutch) fry and fingerlings that were caused by masses of bacterial rods, now known as F. psychrophilum. The bacterium underwent multiple taxonomic reappraisals and was eventually placed within the genus Flavobacterium


17 Francisella noatunensis



Francisella noatunensis

Esteban Soto1* and John P. Hawke2


Department of Medicine and Epidemiology, School of Veterinary Medicine,

University of California Davis, Davis, California, USA; 2Department of

Pathobiological Sciences, School of Veterinary Medicine, Louisiana State

University, Baton Rouge, Louisiana, USA

17.1  Introduction

After Piscirickettsia salmonis, which was the first

Rickettsia-like organism that was demonstrated to cause disease in fish (Fryer et  al., 1992), other

Rickettsia-like organisms were reported in Nile tilapia (Oreochromis niloticus) (Chern and Chao,

1993; Chen et  al., 1994), blue-eyed plecostomus

(Panaque suttoni) (Eigenmann & Eigenmann) and the grouper Epinephelus melanostigma (Khoo et al., 1995; Chen et al., 2000; Mauel et al., 2003).

Kamaishi et al. (2005) amplified and sequenced the

16S rDNA from an agent in frozen kidneys from diseased three-line grunt (Parapristipoma trilineatum) from aquaculture farms. The sequence aligned with other eubacterial 16S rDNA sequences with


18 Mycobacterium spp.



Mycobacterium spp.

David T. Gauthier1* and Martha W. Rhodes2


Department of Biological Sciences, Old Dominion University, Norfolk, Virginia,

USA; 2Department of Aquatic Health Sciences, Virginia Institute of Marine

­Science, The College of William and Mary, Gloucester Point, Virginia, USA

18.1  Introduction

18.1.1  Mycobacterium spp.

Members of the genus Mycobacterium (Order

Actinomycetales, Family Mycobacteriaceae) are aerobic to microaerophilic, non-motile, rod-shaped bacteria that stain Gram positive and acid fast.

Excepting non-culturable species (e.g. M. leprae), mycobacteria are frequently grouped by the phenotypic characters of growth rate and pigmentation

(Runyon, 1959). Runyon Groups I–III are fastidious and take more than 5 days to produce colonies on solid media. Group I mycobacteria are photochromogenic, producing yellow–orange pigment (Fig. 18.1) on exposure to light, and include species such as


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