Aquaculture Virology

†Deceased.

Preface

Frederick S.B. Kibenge and Marcos G. Godoy, EDITORS

The textbook Aquaculture Virology covers the principles of clinical virology for viruses of the three major aquatic food animals: fish, crustaceans and mollusks. The increasing importance of these aquatic animals as an excellent source of quality protein for human consumption has resulted in increased interest in aquatic animal viral diseases, which are the principal natural limiting factors for aquaculture worldwide, causing massive losses in farmed and occasionally in wild populations. Due to the rarity of effective viral vaccines, and in the absence of antiviral drugs for use in aquaculture, prevention and effective control of viral diseases rely very heavily on biosecurity, hygiene and attempted eradication by depopulation. Such control measures depend on an understanding of all aspects of the virus and its relationship to the disease and epidemiology, including interaction between wild and farmed populations. This is an area that has desperately needed a comprehensive textbook for at least two decades; this book attempts to fill that knowledge gap and become the go-to book for this topic.

To date, textbooks on viruses infecting fish, crustaceans and mollusks have been on the whole disease-centric and focus on individual viral diseases selected based on epizoo-centric approaches, with little to no coverage of the basic biology of the viruses. This is in contrast to textbooks on viruses infecting terrestrial—farmed, pet and free-range (wild)—animals, as well as humans. Where aquatic animal (mostly fish) viruses are included in the various animal virology textbooks, they are consistently relegated to the last pages of the various chapters, in most cases with very limited information. Moreover, there is no context for aquaculture being described in a similar way as the terrestrial viruses impacting agriculture or human health. Despite considerable advances in animal virology in recent years, coupled with an economically important global aquaculture industry, knowledge of viruses of animal aquaculture is still sparse and in some cases outdated, although these viruses are closely related to well-known virus families.

The last book in fish virology (Fish viruses and fish viral diseases, 1988, Wolf, K.) was published in the 1980s. A lot of work has been done on fish viruses, and many new aquatic animal viruses continue to be discovered. The editors of Aquaculture Virology wanted to present the current state of knowledge of aquatic animal viruses within the current virus classification and taxonomic context, thereby allowing readers to draw on the principles of general virology.

The authors were encouraged to use the same format in all chapters to facilitate the reading and studying. Thus whereas each chapter includes the main specific diseases that affect the aquatic organism, the description of each disease includes disease name, structure and composition of virus, classification and virus replication, epidemiology, pathology and immunity, diagnostic methods (gross pathology, histopathology, cell culture, polymerase chain reaction, sequencing, enzyme-linked immunosorbent assay, etc.) and prevention and control. The information in the book provides the reader with readily applicable and updated knowledge, delivered in a systematic and succinct way. Those interested in aquatic animal viral diseases will find that the book provides comprehensive, high-quality information in a single source, detailing the diseases, viruses, pathology, laboratory methods for the detection and confirmation of the virus and the prevention and control of these diseases within aquaculture systems. This book targets a unique audience knowledgeable in aquaculture and fisheries with interest in animal pathogens, specifically viruses. This book will be used by researchers, teachers, students, diagnostic laboratory staff, clinical veterinarians, aquaculture disease practitioners, farmers and all people interested in viruses in general.

The authors for the different chapters are invited international experts recognized in their specific virus family or diseases. The editors gratefully acknowledge all the authors for their contributions and keeping to the deadlines.

Part I

General Aspects

Outline

Chapter 1 Introduction to Aquaculture and Fisheries

Chapter 2 Classification and Identification of Aquatic Animal Viruses

Chapter 3 Unclassified and Unassigned Aquatic Animal Viruses

Chapter 4 Diagnosis of Aquatic Animal Viral Diseases

Chapter 5 Prevention and Control of Viral Diseases in Aquaculture

Chapter 6 Determinants of Emergence of Viral Diseases in Aquaculture

Chapter 1

Introduction to Aquaculture and Fisheries

F.S.B. Kibenge,    University of Prince Edward Island, Charlottetown, PE, Canada

Abstract

Farmed fish, crustaceans and mollusks provide an increasing fraction of the human food supply, supplementing and substituting the overfished or declining wild-catch fisheries, and are of major economic importance in many countries. As in the case of terrestrial agriculture, bringing together large numbers of animals of a single species (ie, monoculture) increases the risk of infectious disease outbreaks, including viral infections. Aquaculture, where farmed fish, crustaceans and mollusks are kept at high population densities in close proximity with wild fishery, is ideal for the emergence of wild-type pathogens that exist benignly in local aquatic ecosystems. This introductory chapter describes the structure of the global aquaculture industry (fish, crustaceans, mollusks, in cold water and warm water, commercial and noncommercial, small and big operations, etc.) and the role of fisheries in supplying protein food to the growing human population. This discussion will help to put the virology in the following chapters in context.

Keywords

Fish; crustaceans; mollusks; aquaculture; fisheries

1.1 Introduction to Aquaculture and Fisheries

The Food and Agricultural Organization of the United Nations (FAO) attributes aquatic organisms that are harvested by an individual or corporate body that has owned them throughout their rearing period to aquaculture, while aquatic organisms that are exploitable by the public as a common property resource, with or without appropriate licenses, are harvested by fisheries (FAO, 2015). The contribution of wild-catch fisheries as a protein source for human consumption plateaued in the mid-1980s, and since then, most natural stocks in marine waters have been harvested at or near maximum rates. According to the FAO, aquaculture remains one of the fastest-growing animal food-producing sectors worldwide. In fact, it is the only animal food-producing sector growing faster than the human population, and it provides an acceptable supplement to and substitute for wild fish. In fact, in 2014, aquaculture contributed more to aquatic animal food destined for human consumption than fisheries for the first time. This share of aquaculture is projected to rise to 62% by 2030, as catches from wild capture fisheries level off (FAO, 2014). Fish, crustaceans and mollusks represent the most economically important global aquaculture industry subsectors (Fig. 1.1), at 66.6 million tonnes in 2012, with an estimated total farmgate value of US$144.4 billion (FAO, 2014), and they are expected to reach US$202.96 billion by 2020 (Grand View Research Inc, 2014).

Figure 1.1 World aquaculture by farmed species sector, 2012. (A) Aquaculture industry subsectors by quantity 2012; (B) aquaculture industry subsectors by value. Developed from FAO, 2014. The State of World Fisheries and Aquaculture 2014. FAO, Rome, 223 pp.

Most of the global aquaculture production (89% by volume) is located in countries in Asia (the most important being China, India, Vietnam and Indonesia). Table 1.1 summarizes the distribution of the three main aquaculture industry subsectors among the top 15 countries that account for 92.7% in global aquaculture production.

Table 1.1

Distribution of the Aquaculture Industry Subsectors Among the Top 15 Countries in Global Aquaculture Production in 2012a

aDeveloped from FAO, 2014. The State of World Fisheries and Aquaculture 2014. FAO, Rome, 223 pp.

bSome of the production data are not available (marine aquaculture), or the production volume is regarded as negligibly low (NL).

1.2 Aquatic Species in Aquaculture

Among the three main aquaculture groups (fish, crustaceans and mollusks), there were 354 fish species, 59 crustacean species and 102 mollusk species recorded by FAO as cultured in 2012. Carp dominates production in both China and the rest of Asia; for Europe and South America, it is salmonids; African aquaculture production is almost exclusively tilapias; and for Oceania, shrimps and prawns dominate (Hall et al., 2011). The most significant farmed species, with more than 100,000 tonnes apiece in 2012 (FAO, 2012), are listed in Table 1.2. However, analysis of aquaculture production and details about farmed species remain only an approximation because many indigenous aquatic species are used in aquaculture without being registered individually in national statistics. For example, in China, more than 200 species are farmed commercially, but total production is registered under fewer than 90 species and species groups. Similarly, in India and Vietnam, the number of cultured species far exceeds the number included in statistics (FAO, 2014). Moreover, for the most widely farmed species, tilapia, the true number of producer countries is higher than the FAO tally of 135 countries because commercially farmed tilapia is not yet reflected separately in national statistics in Canada and some European countries (FAO, 2014).

Table 1.2

Major Farmed Species

FW, freshwater; D, diadromous (these fish spawn in freshwater and migrate to the sea for the main growing period); M, marine.

aMajor farmed species or species group (with production of 100,000 tonnes or more in 2012). Data from FAO, 2012: The State of World Fisheries and Aquaculture 2012. Food and Agriculture Organization of the United Nations, Rome.

1.3 Aquaculture Techniques, Systems and Facilities

The techniques, systems and facilities used in aquaculture are as diverse as the species currently being raised and also vary by geographical region (Bostock et al., 2010), but normally encompass three stages: incubation/hatchery seed production, early rearing and on-growing (fish aquaculture briefly reviewed by Roberts and Shepherd, 1974). In salmon and trout aquaculture, on-growing has merged with husbandry breeding (Aarset and Borgen, 2015). For new farmed species like cod, two methods are mainly used: one is based on capturing wild cod for on-growing, while the other focuses on the production of cod from hatchery seed to market size. Currently, on-growing of wild cod is more economically efficient than using farmed juveniles (FAO, 2016). For a few species, such as eels (Anguilla spp.), farming still relies entirely on wild seed. Although the life cycle of the Japanese eel (Anguilla japonica) in captivity has been completed, techniques for mass production of glass eels have not yet been established because of various technical difficulties (Masuda et al., 2012; Bird, 2013).

World aquaculture production takes place in a number of locations: on land (using freshwater or saline water); in static water ponds (eg, for growing channel catfish and other warm-water species); concrete raceway systems; freshwater gravity tanks; large recirculation tanks; in floating net cages in a lake or river (eg, for growing salmonids and other cold-water species that require clean water with high oxygen levels); offshore (mariculture or marine aquaculture) in floating net cages in the sea and intertidal zones (Shepherd, 1993); in coastal lagoons or brackish-water ponds; and in onshore-based tanks using pumped seawater (eg, for flat fish farming of turbot or halibut that normally lie on the bottom and will not shoal within the water column inside a floating cage (Shepherd, 1993)).

1.3.1 Fish Aquaculture

The top 10 cultured fish species are carps, except tilapia, Pangasius catfish and Atlantic salmon (Table 1.2). China, the largest producer of farmed seafood (61.7% share of the world total in 2012; Table 1.1), grows carp mostly in ponds and rice paddies, with little or no effort to actively nurture the animals, and has been practicing this type of aquaculture for thousands of years. Most carp farming is pond-based, with several species stocked in the same pond (polyculture) (SPC, 2011). Tilapia, and sometimes catfish, are stocked along with carp. Single-species culture (monoculture) is rare, except in flow-through systems and cage cultures of common carp in streams or canals. There are different stocking models for polyculture, depending on the availability of the main source of feed. If grasses (aquatic or terrestrial) are abundant, grass carp can be stocked as the major species. The leftover feed and grass carp excreta would sufficiently fertilize the pond water for the growth of filter feeders. Pond-based carp culture has been traditionally integrated with crop farming (mulberry, fruit, vegetables, etc.) and animal husbandry (ducks, swine, chicken, etc.) in China. The practice has been widely introduced to many other parts of the world, with some modifications to suit local conditions (SPC, 2011).

In industrial aquaculture, farmed fish are reared at high population densities in floating open-net cages near shore (for marine aquaculture), or in a lake or river (for inland aquaculture), in the same water column as the wild fish reservoir, which is often at a relatively low population density (Hill, 2005). The cages are either square/hexagonal steel platform constructions linked together or 10–50-m-deep circular polyethylene rings with netting to hold the farmed fish. Expansion can simply be by cage volume or number of cages. Such a cozy aquatic environmental interaction is inevitably ideal for selecting and propagating more virulent variants of wild-type pathogens that exist benignly in local wild stocks. This so-called local effect is an inherent and highly costly risk of aquaculture throughout the world (Hill, 2005), although for several viral diseases of economic importance to aquaculture, there is no clear evidence for transmission of viruses between wild and farmed fish (Johansen et al., 2011).

Today, in various countries, wild fish are collected from rivers for raising juvenile fish in hatcheries and then released for the restocking of natural habitat and under enhancement programs for recreational fishing purposes, which helps to develop large fisheries in some rivers and inland lakes (Dexter and O’Neal, 2004).

1.3.2 Crustacean Aquaculture

The most recent dramatic change in aquaculture has been the explosive growth in shrimp farming in Southeast Asia (Cressey, 2009). Production has increased almost exponentially since the mid-1970s. This rapid increase largely reflects the dramatic increase in white leg shrimp (Penaeus vannamei) culture in Asia (eg, China, Thailand and Indonesia) and Latin America (Ecuador and Mexico) (Bondad-Reantaso et al., 2012), where the penaeid shrimps have tended to dominate due to high-value and short production cycles and accessible technologies (Bostock et al., 2010). Crustacean farming, similar to marine fish aquaculture, requires a high-quality diet usually containing fish meal and often fish oil as well (Bostock et al., 2010).

1.3.3 Mollusk Aquaculture

For the aquaculture of mollusks (oysters, clams, scallops, abalone, mussels, cockles and related species), there are many varieties of culture and grow-out methods that are particular to specific species and locations (Elston, 1999; Elston and Ford, 2011; Dumbauld et al., 2009). Most mollusk culture requires no feed input (algae are used to feed the larvae and juveniles in hatcheries to provide spat for mollusk culturing). Seed bivalves are produced either in land-based hatcheries or from natural populations, and can be transported over considerable distances to grow-out sites in estuaries, and even to different countries (FAO, 2013). Farming mollusks is limited in several countries because of the limited capacity to produce seed from mollusk hatcheries and nurseries (FAO, 2014), relying on wild seed. The use of wild seed is impossible if the natural populations of mussels are not suited to culture. Intensification will require a reliable, plentiful and inexpensive supply of hatchery seed, requiring the selection and management of broodstocks and the controlled production of high-quality seedstocks (Elston and Ford, 2011).

1.4 Aquaculture Pathogens

The most common cause of infectious diseases in aquaculture is bacteria (54.9%), followed by viruses (22.6%), parasites (19.4%) and fungi (3.1%) (reviewed in Kibenge et al., 2012). Viral diseases have been more difficult to control due to the high susceptibility of aquatic animals to them at an early age, limited availability of therapeutics, insufficient knowledge of pathogenesis of virus infections and limited knowledge about natural resistance mechanisms in aquatic animals. Thus, viruses are the principal pathogens that are negatively affecting aquaculture worldwide.

1.5 Aquaculture Health Management

The intensive rearing of fish, crustaceans and mollusks is highly vulnerable to infectious diseases, similar to captive livestock agriculture (eg, the poultry and swine industries). However, in contrast to terrestrial farmed animals, the animal strains used in aquaculture usually have been derived from wild strains in recent years (Stuart et al., 2007; Duarte et al., 2007; Rosenlund and Skretting, 2006; Svasand et al., 2004) and may not have had enough time to adapt to high-density confinement within the aquaculture environment (Rodriguez-Ramilo et al., 2011). This chronic stress (Snieszko, 1974; Wedemeyer, 1996; Weyts et al., 1999; Yada and Nakanishi, 2002) provides opportunities for the emergence of diseases caused by pathogens that may have been harmless under natural conditions. Several strategies that are used to successfully rear terrestrial farm animals in intensive production systems are also used to prevent and control viral diseases in aquaculture, including the selective breeding for increased resistance against diseases (Gjedrem, 2010) and vaccination (Gudding, 2014). However, despite the traditional use of antibiotics, there is no field experience with applying antiviral drug treatment to farmed fish, although prophylactic antiviral compounds, if available, would be best used in broodstock and valuable genetic stocks that are not used for human consumption (Kibenge et al., 2012). Vaccination is not an option with farmed crustaceans and mollusks because of their primitive immune systems. Thus, reducing the risk of disease in these cases will largely depend on the application of better husbandry and biosecurity management practices at the farm level and the use of quality, high-health, specific pathogen-free seed supported by strict biosecurity (Bondad-Reantaso et al., 2012). Additional biotechnology efforts are being directed at better understanding of immune priming in invertebrates and developing innovative antiviral drug treatment, such as by RNA interference (Rowley and Pope, 2012).

References

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Chapter 2

Classification and Identification of Aquatic Animal Viruses

F.S.B. Kibenge,    University of Prince Edward Island, Charlottetown, PE, Canada

Abstract

The classification of microorganisms is now considered mature. There are over 1000 individual viruses that cause infections of veterinary significance in the major animal species/groups; most are assigned to virus families. Despite considerable advances in animal virology in recent years, coupled with an economically important global aquaculture industry, knowledge of viruses of animal aquaculture is still sparse and in some cases outdated, although these viruses are closely related to well-known virus families. The purpose of this chapter is that viruses within groups share characteristics that help in our understanding of the epidemiology and pathogenesis of infections they produced. If you know the member viruses of a particular family and understand the group properties of each family, it is possible to know some vocabulary about the particular viral disease. Furthermore, this knowledge gives valuable information on laboratory methods used to properly identify new viruses.

Keywords

Virus family; virus genus; virus species; virus genome; virus morphology; International Committee on Taxonomy of Viruses (ICTV); Baltimore classification; virus replication; virus pathogenesis; virus isolation; virus detection; molecular detection; nucleic acid sequencing

2.1 Viruses Are Entities at the Edge of Life

Viruses occupy a unique position in biology. Although they possess some of the properties of living systems (such as a genome), they are in fact nonliving infectious entities and should not be considered microorganisms. Viruses have two clearly defined phases in their life cycle. A virus particle outside a host cell is metabolically inert (functionally inactive). Inside their host cell, the viruses are metabolically active (functionally active); this is their replicative phase, in which the viral genome exploits the machinery of the host cell to produce progeny genome copies, viral messenger RNA and viral proteins that assemble to form new virus particles. Structural features of a virus are determined by requirements for virus multiplication in a host cell (ie, assembly, release, transmission, attachment, penetration and uncoating).

2.2 History of Virus Classification

Schemes for virus classification have changed rapidly in the past few years and continue to evolve. Schemes used in the past included those based on common organ tropisms and common ecological and transmission characteristics of the virus infection, leading to names such as enteric viruses, respiratory viruses, viscerotropic viruses, arboviruses (arthropod-borne viruses), etc. While these systems were useful (and are still useful when discussing diagnosis, pathogenesis and epidemiology), it quickly became clear that groups of viruses affecting the same organ in animals, or with similar epidemiological features, actually consisted of a diverse array of quite different viruses. Subsequent taxonomic systems have focused on the viruses themselves, and the advances in classification have generally followed advances in our knowledge of virus morphology, genomic composition and replication strategy.

2.3 Morphology and Composition of Animal Viruses

A virus particle (or virion) is a structure that has evolved to transfer nucleic acid from one cell to another (which is the function of the virion). The central structure of every virus is a nucleic acid (genome) surrounded intimately by a protein shell or coat (capsid). Together, they form a nucleocapsid.

Genetic economy dictates that the capsid/nucleocapsid structure be built from identical copies of a small number of viral proteins. These protein molecules are arranged to provide maximal contact and noncovalent bonding among subunits (polypeptides) and structural units. The repetition of such interactions among a limited number of proteins results in a regular structure of either icosahedral or helical symmetry. Thus during virus multiplication in an infected cell, viral structural proteins associate spontaneously (through random thermal movement and are bonded in place through weak chemical bonds) to form capsomeres. The capsomeres self-assemble in a defined manner, giving the capsid a characteristic symmetry (icosahedral or helical) (Fig. 2.1A). Viruses that do not fall into either category are referred to as being complex (eg, poxviruses and bacteriophages).

Figure 2.1 General structure of viral capsids. (A) Spontaneous association of viral structural proteins (VPs) and self-assembly of capsomeres into capsids with characteristic symmetry (icosahedral or helical); (B) Icosahedral symmetry with a 5:3:2-fold rotational symmetry; (C) Various clustering of structural units (trimers, pentamers, hexamers or dimers).

Virus capsids of icosahedral symmetry have 12 corners, 20 equilateral triangular faces and 30 edges with a 5:3:2-fold rotational symmetry, which passes through their corners, faces and edges, respectively (Fig. 2.1B). This symmetry is the most efficient of possible arrangements for repeating subunits in a closed shell, in the sense that it uses the smallest unit to build a strong structure enclosing a maximum volume. Thus, in the surface of any icosahedrally symmetrical structure, there are exactly 60 identical elements (and certain multiples of 60) related to each other by twofold, threefold and fivefold rotation axes. Viruses generally fit 60 × N subunits into their capsids. N is sometimes called the triangulation number, also abbreviated T. Various clustering of structural units (trimers, pentamers, hexamers or dimers; Fig. 2.1C) give the characteristic appearance of capsomeres in electron microscopy (EM) and may be used to identify a virus. Viewed by EM with negative staining, icosahedral virions appear hexagonal or spherical in outline.

Virus capsids of helical symmetry have a single rotational symmetry. The length, width and pitch of the helix (ie, the distance covered by each complete turn of the helix), as well as the number of capsomeres in each turn of the helix, are constant for a given virus type. Viewed by EM with negative staining, helical virions appear pleomorphic (if they lack a matrix protein in the envelope) but in native state, they are spherical (except rhabdoviruses, which are rod- or bullet-shaped, and bafiniviruses and roniviruses, which are bacilliform).

Some viruses are unenveloped (or naked), and exist basically as nucleocapsids. Many other animal viruses acquire an envelope by passing through one of the cell membranes (nuclear membrane, plasma membrane, rough endoplasmic reticulum, Golgi complex). Thus, the actual content of a viral envelope depends on the host cell and the cellular membrane through which the particle buds, but is generally comprised of a lipid bilayer with transmembrane viral glycoproteins. In viral envelopes, the protein is viral coded and the carbohydrate is added by cellular glycosyl transferases. The glycoproteins are the surface structures (spikes, peplomers) on enveloped viruses; they therefore form the viral attachment proteins essential for both host specificity and viral infectivity, and epitopes targeted for virus neutralizing antibodies, as well as for the effector cells of cell-mediated immunity. The viral matrix protein (nonglycosylated envelope protein) adds rigidity to the enveloped virions. Enveloped viruses are ether-sensitive and chloroform-sensitive (ie, infectivity of enveloped viruses is destroyed by detergents and lipid solvents), which is important when determining the morphology of a virus.

The morphology of a virus particle includes the size (ranging from 12 to 430 nm in diameter) and shape of the virion (spherical, rod-shaped, ovoid, bacilliform, bullet-shaped, pleomorphic), number of capsomeres, symmetry of capsid (icosahedral, helical, complex) and presence or absence of an envelope. This is the primary criterion for virus classification (see also Fig. 2.4). The viral genome can be either RNA (and nonsegmented or segmented) or DNA (and linear or circular) and single-stranded (positive sense or negative sense or ambisense) or double-stranded (see also Tables 2.2 and 2.3).

Regardless of the structural complexity of a virion, proteins are its principal constituent. The viral proteins can be structural or nonstructural proteins. The structural viral proteins include surface proteins that make up the capsid and viral enzymes like neuraminidase, and internal proteins such as the viral enzymes DNA polymerase, RNA polymerase, proteases, helicases and ligases.

The surface viral proteins (viral attachment proteins) have an affinity for complementary receptors on the surface of susceptible cells and contain antigenic determinants (epitopes), which is important for vaccines as immunogens, for virus identification (serotypes) and for diagnosis (serology). Nonstructural proteins are not found in the virion; rather, they are only present in infected cells during virus replication and are involved in virion assembly. Such nonstructural proteins have been used to develop serological tests that differentiate between infected and vaccinated animals (DIVA).

2.4 Overview of Virus Replication

Virus replication occurs only in living cells, in sequential steps involving the attachment, penetration, replication of the parent virus and release of progeny viruses. Every animal virus family (refer to Tables 2.2 and 2.3) has a different strategy of replication (a useful parameter in virus classification). The strategy of virus replication depends on the characteristics of the virus genome and the pathway they use to produce their messenger RNA (mRNA) (ie, the Baltimore classification; see Section 2.7). The ability of the virus to multiply and the fate of an infected cell depend on the synthesis and function of the viral proteins. These proteins replicate viral genomes, package the genomes into capsids, alter the structure of infected cells and alter the function of infected cells to favor virus replication.

One-step growth curve is a common experimental procedure for studying virus replication. One-step conditions exist when all cells in a culture are infected simultaneously using a high multiplicity of infection (MOI; ie, the ratio of virus particles to cells) so as to prevent secondary cycles of infection. After the initial infection, the excess unabsorbed virus is rinsed off, and the amount of infectious virus over time is followed by sequential sampling and titration. When these titers are plotted as a function of time, a growth curve of a given virus in a given cell system is generated. A virus that is free in the cell culture medium can be titrated separately (for the yield of extracellular virus) from a virus that remains cell-associated (the yield of intracellular/cell-associated virus). We recognize four stages of a virus growth curve (Fig. 2.2): attachment, eclipse period (infectious virus particles cannot be demonstrated inside the cell), maturation and release (by cell lysis in naked viruses and by budding in enveloped viruses).

Figure 2.2 A one-step growth curve for an animal virus. (A) One-step growth curve for a naked animal virus; (B) One-step growth curve for an enveloped animal virus budding from the cytoplasmic membrane. Note the differences between intracellular and extracellular virus titers for both the naked virus (A) and the enveloped virus (B). The top bar demarcates the four stages of a virus growth curve: 1, attachment and penetration; 2, eclipse period; 3, maturation; 4, release. In enveloped viruses, stages 3 and 4 occur simultaneously. Stages 2 and 3 make up the latent period.

There are eight essential steps in the virus replication cycle. In order to initiate infection, the virus must bind to the cell surface via capsid or envelope proteins. It must then penetrate the cell and uncoat to make its genome accessible to viral and host machinery for transcription and/or translation, and transport to sites where virus-directed synthesis takes place. If replication occurs in the nucleus, viral proteins have to be transported to the nucleus for virion assembly.

1. Attachment (adsorption) to cell surface: Depends on the presence of specific cell receptors for complementary viral attachment proteins on the surface of the virion (ligands or antireceptors). This specificity determines the tissue tropism and host range of a virus. If a cell does not have receptors for the virus, then that cell and that animal will be resistant to that virus. Adsorption of virus may require calcium ions (to reduce electrostatic repulsion) but does not require energy or temperature.

2. Penetration (uptake): Can be by at least one of four mechanisms; however, some viruses can use more than one mechanism. These processes require energy and are temperature-dependent. Therefore, the cell must be metabolically active for these mechanisms to occur:

 Translocation: The entire virus particle crosses the cell membrane intact. This is relevant only to naked viruses.

 Genome injection: Viral genome penetrates cytoplasm via a pore that has been created in the plasma membrane. This is relevant only to naked viruses.

 Receptor-mediated endocytosis (viroplexis): (a) for naked viruses, this occurs via clathrin-coated pits; (b) for enveloped viruses, this occurs via membrane fusion with the endosome membrane.

 Membrane fusion directly with plasma membrane of the cell. This is relevant only to enveloped viruses.

3. Uncoating: Makes viral genome accessible to viral or cell machinery for transcription or translation.

4. Transcription (viral genome expression): Eukaryotic cells lack enzymes for synthesizing mRNA off a viral RNA genome, either in the nucleus or in the cytoplasm; and they cannot transcribe viral DNA located in the cytoplasm. Therefore, only DNA viruses that replicate in the nucleus utilize the cellular machinery (cellular RNA polymerase II and other cellular enzymes) for transcription. All other viruses provide their own enzymes to produce mRNAs. The classification schemes used for animal viruses (refer to Tables 2.2 and 2.3 and Section 2.7) emphasize the different strategies of viral genome expression.

5. Translation: This is the key event in viral replication. The eukaryotic cell translation apparatus works only on monocistronic mRNA. Polycistronic mRNA is translated into a precursor polyprotein, which requires posttranslational cleavage into individual mature proteins. In DNA viruses, early proteins serve as viral enzymes and regulatory proteins, whereas late proteins are structural proteins. Viral proteins may undergo posttranslation modification (eg, glycosylation) and are then transported to sites where virion assembly occurs.

6. Replication of viral nucleic acid: Like transcription, it differs between virus families.

7 and 8. Assembly, maturation (morphogenesis) and release. These four processes occur with the two types of viruses, as follows:

 Naked viruses: Virus release occurs by cell lysis. Structural proteins of naked icosahedral viruses associate spontaneously to form capsomeres, which self-assemble to form empty procapsids, into which viral nucleic acid is specifically packaged (viral genomes contain a packaging sequence that directs its encapsidation. Virions accumulate within the cytoplasm/nucleus.

 Enveloped viruses: Virus release occurs by budding. Viral envelope proteins are transported and inserted into the cellular membrane at the site where virus buds. In many virus types, the final stage of virus maturation does not occur until after the virus particle is released from the host cell. Budding may not damage the cell, so many such viruses (eg, arenaviruses, retroviruses) are noncytopathogenic and are associated with persistent infections; others are cytolytic (eg, herpesviruses).

2.5 Current Virus Classification Scheme

The International Committee on Taxonomy of Viruses (ICTV) in 1973 established a single, universal taxonomic scheme for all viruses regardless of host, which is set arbitrarily at hierarchical levels of order, family, subfamily, genus and species, for example:

 Order is named with the suffix virales (eg, Nidovirales)

 Family has the suffix viridae (eg, Coronaviridae)

 Subfamily has the suffix virinae (eg, Torovirinae)

 Genus has the suffix virus (eg, Bafinivirus)

 Species are given by a full name (eg, White Bream virus).

The first letters of virus order, family, subfamily and genus terms are capitalized, and the names are printed in italic or underlined.

The ICTV currently lists 2618 virus species. The genomes of about 40,000 viral strains, across more than 900 species, have been sequenced (reviewed by Sharma et al., 2015). Based on the Ninth Report of the International Committee on Taxonomy of Viruses (King et al., 2012), the order is the highest taxon in virus classification. Virus orders represent groupings of families of viruses with good evidence of phylogenetic relationship among them; ie, they share a common evolution (ancestry), as evidenced by common nucleotide sequences between virus families, similar gene arrangements and gene products and similar mechanisms of gene expression, and similar virion morphology.

Virus families represent groupings of genera that share common characteristics and are distinct from the member viruses of other families. This level in the taxonomic hierarchy is stable, and it is the benchmark of the entire taxonomy system (Fig. 2.3). Today, classification of viruses into families is based principally upon four criteria:

 The structure of the virion—that is, size and shape of virion, number of capsomers, symmetry of capsid and presence or absence of envelope.

 Type and character of the viral genome—that is, type of nucleic acid (DNA or RNA), molecular weight, linear or circular, single- or double-stranded segmented or nonsegmented, polarity, nucleotide sequence, etc.

 Strategy of viral replication—this depends on type of genome, etc. Positive-sense single-stranded RNA (ssRNA) viruses behave like mRNA once inside a host cell. Negative-sense ssRNA viruses and double-stranded RNA (dsRNA) viruses have viral transcriptase. DNA viruses multiplying in the nucleus use cellular enzymes, and those multiplying in the cytoplasm carry their own enzymes.

 Possession of characteristic enzymes—for example, reverse transcriptase, neuraminidase, etc.

Figure 2.3 Diagrams of animal viruses arranged by type of nucleic acid and the presence or absence of a viral envelope. (A) Orders, families and subfamilies of viruses that infect invertebrates (including crustaceans and mollusks); (B) Orders, families and subfamilies of viruses that infect vertebrates (including fish). From King, A.M.Q., Adams, M.J., Carstens, E.B., Lefkowitz, E.J., 2012. Virus Taxonomy. Ninth Report of the International Committee on Taxonomy of Viruses. Academic Press, New York, NY. Copyright © Elsevier (2012), with permission.

Some families are so complex that subfamilies have been introduced to break them into manageable divisions (Fig. 2.3).

Table 2.1 summarizes the distribution of the different virus families that occur in the three major aquatic food animals (fish, crustaceans and mollusks), which are discussed in the specific chapters in this book. Interestingly, only one virus family (Birnaviridae) occurs in all three major aquatic animal host species, and given the potential food-web among the aquatic animals, it follows that natural transmission pathways and interspecies (fish, crustaceans and mollusks) reservoirs of many of these viruses are limited. There is still a wide range of viruses that cannot be classified using this scheme (see Chapter 3: Unclassified and Unassigned Aquatic Animal Viruses). Many viruses, particularly those of crustaceans and mollusks, are inadequately identified, poorly described, and attributable sequences are available for only a small number of economically significant viruses, poorly reflecting the diverse nature of viruses of animal aquaculture. In some cases, presumptive viruses that are apparent in histological sections (Bonami et al., 1992; Edgerton et al., 1994) or in ultrathin sections for EM (Dunlap et al., 2013; Granzow et al., 2014) have not been further characterized. Examples of unclassified viruses in aquaculture and fisheries are listed in Section 2.8, later in this chapter. It is considered that the number of unclassified viruses will likely continue to increase as viral genomes with unique architectures are identified through increasing application of metagenomic approaches. Viral metagenomics has already challenged current classification schemes by revealing new viruses that blur the boundaries between groups once thought to have well-defined and unique characteristics (Rosario et al., 2009, 2012; Reuter et al., 2015). On the other hand, there is at least one situation where a disease in mollusks (such as hemic neoplasia of the clam Mya arenaria) that previously was considered to have a viral etiology has been determined not to be the case (Arriagada et al., 2014). The neoplastic hemocytes are transmitted from individual to individual—so the transmissible element appears to be the cancer cell itself, rather than a virus (Metzger et al., 2015).

Table 2.1

DNA and RNA Viruses Infecting the Three Major Aquatic Food Animals (Fish, Crustaceans and Mollusks)

aThe following DNA virus families of vertebrates do not occur in fish species: Herpesviridae, Asfarviridae and Anelloviridae.

bBivalve mollusks are filter feeders and consequently may bioaccumulate in their tissues viruses from humans and other vertebrates (Meyers, 1984). Only families of viruses reported to be infective for bivalves are included in this table, and discussed in the specific chapters. Indeed, Meyers (1980) and Meyers et al. (2009) have shown that the 11-segment American oyster isolate 13p2 does not productively infect oysters and have argued that the putative aquareoviruses obtained from oysters or clams are more likely to be fish viruses that simply accumulated in these shellfish upon filter-feeding of virus-contaminated water (Nibert and Duncan, 2013).

cA full genome sequence of a novel salmon parvovirus—the first parvovirus to be identified in a fish species—was obtained from sockeye salmon smolts from British Columbia–Canada, but it is not known yet if it is associated with disease (Miller et al., 2013).

dA novel Hepadnavirus that infects the white sucker Catostomus commersonii from the Great Lakes region of the United States was recently reported (Hahn et al., 2015).

eA novel iridovirus was reported in a mollusk (Gregory et al., 2006). Three other iridovirus infections have been reported in bivalve mollusks (CEFAS, 2011).

fNudiviridae is a newly approved virus family (Jehle et al., 2013).

gJEECV is a novel DNA virus (with 15-kbp circular genome) isolated from samples showing endothelial cell necrosis in Japanese eel, Anguilla japonica (Mizutani et al., 2011).

hPanulirus argus virus 1 (PaV1), a novel dsDNA virus with morphological features of Herpesviridae and Iridoviridae, infects spiny lobsters (Shields and Behringer, 2004; Montgomery-Fullerton et al., 2007).

iAbalone shriveling syndrome-associated virus (AbSV) belongs to a novel unclassified virus family of bacteriophage-related chimeric marine virus infecting abalone (Zhuang et al., 2010).

jThe following RNA virus families of vertebrates do not occur in fish species: Bunyaviridae, Deltavirus, Filoviridae, Bornaviridae, Nyamiviridae, Arteriviridae, Astroviridae, Flaviviridae and Picobirnaviridae.

kPreviously, there had been several reports of reverse transcriptase activity in diseased clams, suggesting a retrovirus or retrotransposon; and there are also reports that the disease was transmissible, which suggested a virus. Arriagada et al. (2014) used high-throughput sequencing to look for a retroelement and found amplification of a long terminal repeat-retrotransposon, not a retrovirus, associated with disease. Most recently, it has been found that the genotypes of cancer cells in different individuals are nearly identical and do not match their hosts. This shows that the cancer cells are transmitted from individual to individual—so the transmissible element appears to be the cancer cell itself, rather than a virus (Metzger et al., 2015).

lTwo isolates (DF 20/00 and DF 26/02) of novel virus obtained from common carp and koi carp with gill necrosis were ultrastructurally reminiscent of arenaviruses, and may represent the first fish viruses of the family Arenaviridae but may be members of a hitherto undescribed viral taxon (Granzow et al., 2014).

mLSNV is positive ssRNA virus, which on phylogenetic analysis formed a distinct clade between members of Luteoviridae and Togaviridae (Flegel, 2006), and associated with MSGS in shrimp (Sritunyalucksana et al., 2006).

nFisavirus 1 (fish stool-associated RNA virus) is a novel positive ssRNA virus identified from a freshwater carp (Cyprinus carpio) using viral metagenomics; complete genome sequence is 8712 nucleotides long and shares the same genome organization and distant phylogenetic relationship to the currently unclassified posaviruses that may belong to a novel family of the order Picornavirales (Reuter et al., 2015).

Tables 2.2 and 2.3 list the distinguishing features of aquatic food animal virus families based on the Ninth Report of the International Committee on Taxonomy of Viruses (King et al., 2012). This information is bound to change as the discovery of new viruses and the generations of sequence data on older virus isolates are reported. Moreover, new criteria being used by the different ICTV study groups require extensive sequence data. For example, the current taxonomic hierarchy could not recognize phylogenetic grouping of the 11-segment fish reoviruses, grass carp reovirus (GCRV) isolates HZ08 (Wang et al., 2012), GD108 (Ye et al., 2012) and 104 (Fan et al., 2012), and the 10-segment fish reovirus, piscine orthoreovirus (PRV) (Palacios et al., 2010), into a larger, monophyletic taxon (Nibert and Duncan, 2013). Another discrepancy was noted with regard to grouping cutthroat trout virus (CTV) in Hepeviridae (Batts et al., 2011). When complete genome sequences of Hepatitis E virus (HEV) and other HEV-like viruses were compared, CTV fell out of the family group, as it has different positioning of open reading frame 3 (ORF3), resulting in gaps when aligned with other members of Hepeviridae (Oliveira-Filho et al., 2013).

Table 2.2

Distinguishing Viral Properties of DNA Virus Familiesa Containing Veterinary Aquatic Pathogens

N/A, not available.

aSummarized from the Ninth Report of the International Committee on Taxonomy of Viruses (King et al., 2012).

bGenome of DNA virus is invariably nonsegmented; ss, single-stranded; ds, double-stranded. +ve, positive sense; −ve, negative sense. In Parvoviridae, virions may contain a negative or positive sense.

cRT, reverse transcriptase; +#, replicates in cytoplasm.

dAdditional information specific to fish hepadnavirus from Hahn et al. (2015).

eVertebrate iridoviruses are enveloped.

fSummarized from Wang and Jehle (2009). Nudiviridae is a newly approved family (Jehle et al., 2013).

Table 2.3

Distinguishing Viral Properties of RNA Virus Familiesa Containing Veterinary Pathogens

N/A, not available.

aSummarized from the Ninth Report of the International Committee on Taxonomy of Viruses (King et al., 2012).

bAll RNA is linear; 1, nonsegmented; 2–12, number of segments in genome (segmented); ss, single stranded; ds, double stranded; −ve or +ve, negative or positive sense of ssRNA.

cRT, reverse transcriptase.

dAdditional information specific to Tilapia lake virus (TiLV) with 10 genome segments from Bacharach et al. (2016).

eDouble capsid.

2.6 Classification of Aquatic Animal Viruses of Veterinary Importance

Aquatic food animals are cold-blooded (poikilothermic) animals and are either vertebrates (eg, fish) or invertebrates (eg, crustaceans and mollusks). Aquatic animal virology started in the early 1960s using EM morphological data for crustacean viruses (Vago, 1966) and with the establishment of fish cell lines for the isolation of piscine viruses (Wolf and Quimby, 1962; Gravell and Malsberger, 1965). According to the Food and Agricultural Organization of the United Nations (FAO), aquaculture is set to remain one of the fastest-growing animal food-producing sectors in the world. In fact, it is the only sector growing faster than the human population. Fish, crustaceans and mollusks represent the most economically important global aquaculture industry sectors, at 66.6 million tonnes in 2012 with an estimated total farmgate value of US$144.4 billion (FAO, 2012) and expected to reach US$202.96 billion by 2020 (Grand View Research Inc., 2014). Information on viruses of these aquatic animal species of economic importance is lagging behind that of viruses of homeothermic vertebrates (ie, terrestrial viruses affecting agriculture) (Sano, 1995; Essbauer and Ahne, 2001). The increasing demand for seafood around the world has resulted in increased interest in aquatic viral diseases, which are the principal natural limiting factors for fish aquaculture (Kibenge et al., 2012a) and shrimp aquaculture (Flegel et al., 2008; Johnson et al., 2008; Stentiford et al., 2012) worldwide, causing massive losses in farmed and wild populations.

Improved diagnostic and surveillance efforts, which have accompanied the burgeoning aquaculture industry, have been accompanied by the discovery of new and emerging viral diseases that are naturally endemic in wild aquatic animal populations. Often, new viruses have been discovered only after a serious disease has occurred, although proof of etiology (fulfilling Koch’s postulates of disease causation) and study of pathogenesis are often lacking because the viruses have not yet been cultured. The development of high-throughput sequencing methodology (or next-generation sequencing) (Shendure and Ji, 2008; Metzker, 2010; Munroe and Harris, 2010) and the enabling of high-throughput metagenomics to commercially important fisheries and aquaculture samples have resulted in the discovery of unknown viruses in newly emerging diseases (Batts et al., 2011, 2012; Haugland et al., 2011; Lőrincz et al., 2011; Murray and Peeler, 2005; Palacios et al., 2010; Walker and Winton, 2010; Alavandi and Poornima, 2012; Fichtner et al., 2013; Gjessing et al., 2015; Hahn et al., 2015; Reuter et al., 2015; Bacharach et al., 2016), albeit the taxonomic position and pathogenicity of some of these new viruses remain unresolved (Suttle, 2005, 2007; Sritunyalucksana et al., 2006; Zhuang et al., 2010; Ng et al., 2013; Dunlap et al., 2013).

With the improved bioinformatics tools for nucleic acid sequence analysis and sophisticated databases in virus research (ie, a new field called viroinformatics (Sharma et al., 2015)), it is now necessary in most cases only to obtain nucleotide sequence data on a few viral genes in order to place an unknown virus in its proper taxon (Benkő et al., 2002; Palacios et al., 2010; Lőrincz et al., 2011; Batts et al., 2011, 2012; Haugland et al., 2011), and from there to identify it specifically. The practical result of the current virus classification scheme discussed previously is a system of groups of viruses that have common biological properties, as given in Table 2.4 for aquatic animal viruses of veterinary importance. Viruses within groups share characteristics that help in our understanding of the epidemiology and pathogenesis of infections they produced. If you know the member viruses of a particular family and understand the group properties of each family, it is possible to obtain some vocabulary about the particular viral disease. The information presented in Table 2.4 is bound to change as the discovery of new viruses and the generation of sequence data on older virus isolates are reported. Recently, a novel fish hepadnavirus associated with hepatic neoplasia in the white sucker Catostomus commersonii from the Great Lakes region of the United States was characterized (Hahn et al., 2015). Even more recently, a novel orthomyxo-like virus named Tilapia lake virus (TiLV) causing mass die-offs of tilapia in Israel and Ecuador has been characterized; the virus as 10 genome segments of negative-sense ssRNA, with 5ʹ and 3ʹ noncoding regions characteristic of influenza viruses, and will ultimately be classified as representing a new genus in the family Orthomyxoviridae (Bacharach et al., 2016).

Table 2.4

Major Taxonomic Virus Groups (and Approved/Tentative Species) of Veterinary Importance in Aquaculture