1. health and fitness
2. disease

International Journal of Microbiology and Allied Sciences
November 2014, 1(2):46-77
ISSN: 2382-5537
REVIEW ARTICLE
Page: 46-77
Bacterial and viral diseases in fish & shrimps
Sukhdev Singh*, Manu Bala, Akanksha, Shamsher Singh Kanwar
Department of Biotechnology, Himachal Pradesh University, Summer Hill, Shimla-171005, India.
*Corresponding author. Email address: [email protected]
ABSTRACT
Aquaculture play central role in socio-economic development of rural and coastal countries. Human
interfere displace aquatic animals from their natural ecosystems to artificial stressed environment which
often results in emergence & spread of pathogens and new diseases. Many severe pathogens have been
emerged in aquaculture important species; especially the viral ones. Some economically important
species are Atlantic salmon (Salmo salar), Rainbow trout (Oncorhynchus mykiss), Sea bream (Pagrus
major), Yellow tail (Seriola quinqueradiata), Gilthead sea bream (Sparus aurata), Ayu (Plecoglossus
altivelis), Sea bass (Dicentrarchus labrax), Turbot (Scophthalmus maximus), Flounder (Paralichthys
olivaceus) and shrimps (Penaeus monodon & P. vannamei). Permanent eradication of these pathogens
from major aquaculture producing countries is very necessary; for which the causative etiological agent
must be known. So main aim of this article is to describe these pathogens and developed diseases.
Permanent vaccines against all pathogens are not developed yet as they change their pathogenicity via
enhanced virulence. Additionally, available vaccines are unable to raise strong and long term immune
response. Amongst these pathogens, viral ones are discussed in detail as they are most contagious and
can wipe out whole farms.
Key words: Aquaculture, pathogens, diseases, vaccines, viral pathogens.
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INTRODUCTION
Fish farming is an ancient practice known to human civilization [1]. From 20th century, husbandries
were started to establish for salmon, trout and shrimp species, and thereafter aquaculture became
a major animal food producing sector [2]. Aquaculture species are farmed in fresh, brackish and
marine water environments in which they are reared for optimal production [3]. These species are
diverse in geographically mean, viz. from tropical to arctic & subarctic, from lakes & rivers to
estuaries. Principal farming species are salmon, trout, carp, oyster and shrimps. Countries like UK,
Canada, Norway, Chile and USA are dominant producers of salmon and trouts whereas Asian
countries are involved for carp, oyster and shrimps [4]. In previous decades many serious diseases
have been emerged in aquatic animals due to anthropogenic global movement of animals & their
products. These diseases are listed as ‘notifiable’ by world organization of animal health. Among
the fish pathogens, diseases caused by viruses are of great concern as they wipe out whole
aquaculture farms. Brood stocks from wild fisheries are significant healthy carrier of viral
pathogens [5] and attempts are in progress to confront these carriers using RNA interference
approach [6]. Fish cell lines are suitable subjects for modern fish pathology especially for virology
to isolate the viral agent in vitro [7]. Bacteria also cause severe diseases in fish and economic losses
in aquaculture (Table 1). Some of the bacterial pathogens among these are known to infect (via
food or contact) humans also [8]. Death outbreaks associated with usually occur due to
consumption of raw or improperly processed infected fish. Many bacterial pathogens have been
reported for histamine poisoning which occur by decarboxylation of histidine [9]. These histamine
formers grow optimally at 25 oC. Symptoms of histamine poisoning are diarrhea, dyspnoea,
pruritus, hypertension and metallic or peppery taste in mouth [9]. Like fishes, shrimps also cover
a major part of aquaculture industry and as demanding sea food. Shrimps comprise 15-20% of total
fishery products worldwide [10]. Out of various species, two principal species namely Black tiger
shrimp (Penaeus monodon) and White pacific shrimp (Penaeus vannamei) dominate the shrimp
production as animal food. Pathogens emerge in shrimps also, especially viruses (Table 2);
producing mass mortality in the farm [11]. Some immunity components have been described in
shrimps like toll like receptors [12], RNAi [13] and vaccination attempts are also in progress [14].
So in this review we have summarized the diseases in the light of expanding geographical host
range, virulence diversity, clinical signs and vaccination schemes and emphasized on the viral
pathogens. Complete diagnosis of these pathogens can’t be fully described as they require
histopathological, immunological & molecular diagnostics, in situ hybridization and cell culture
diagnosis [15].
Bacterial diseases in fish
Diseases due to Aeromonads
Aeromonads are the most common bacterial pathogens in fishes throughout the world. Species
included in this group are Aeromonas hydrophila, A. veronii and A. salmonicida which cause
serious diseases in aquatic animals and in humans too [16]. Among these A. hydrophila causes
motile aeromonas septicemia (MAS) or red sore disease [17] and A. salmonicida causes
furunculosis [18]. The most common clinical signs are hemorrhages in skin, fins, oral cavity and
muscles with superficial ulceration of epidermis [19]. Exophthalmia, ascites, swollen kidneys,
splenomegaly, necrosis in spleen and kidneys are also observed. A. salmonicida is antigenically
homogenous and can be diagnosed by immunodiagnostics and PCR [20]. Expression profiling and
virulence factors have been described in A. salmonicida species [21,22]. Bacteriovorax sp. has
recently been identified as biocontrol against A. veronii [23,24].
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Bacterial kidney disease (BKD)
Aetiological agent for BKD or Overt disease is Renibacterium salmoninarum which mainly affect
salmonids [25]. Infection symptoms are exophthalmia, petechial hemorrhages, enlargement &
necrosis in kidneys with granulomatous lesions [26]. Biochemically and antigenically R.
salmoninarum is homogenous [27] and PCR, IFAT and ELISA diagnostic methods have been
developed against the pathogen [28,29]. This pathogen spread horizontally or vertically [30,31]
and transmission dynamics & phylogenetic history have been recently demonstrated with single
nucleotide polymorphism (SNP) analysis [25].
Mycobacteriosis (Fish Tuberculosis)
Fish are convincing host of mycobacteriosis [32]. Many mycobacterium species (M. marinum, M.
fortuitum, M. chelonae, M. avium, M. gordonae, M. abscesus, M. aurum, M. parafortuitum, M.
poriferae and M. triplex) have been recognized as causative agents for this chronic disease [33]
but M. marinum is notable amongst these. Its host range is salmons, turbot, sea bass, striped bass,
sod and halibut [34]. Disease symptoms include exophthalmoses, keratitis, change in pigmentation,
skin ulcerations, granulomas in liver, spleen & kidney and hemorrhagic lesions in muscles [35].
Histopathology and PCR detection procedures have been developed to detect the pathogen [36].
Vibriosis
Pathogenic species in this group are Vibrio anguillarum, V. ordalii, V. vulnificus Biotype 2, V.
harveyii and Alivibrio salmonicida within the family Vibrionaceae [37]. V. anguillarum is
antigenically heterogeneous and approximate 20-25 ‘O’ serotypes has been identified in contrast
to other Vibrio species [38]. Among these only O1, O2 and O3 serotypes are implicated in mortality
cases. Complete genome of V. anguillarum has been sequenced recently [39]. Clinical signs
observed with vibriosis are hemorrhagic septicemia on the base of fins, edematous lesions, corneal
opacity and exophthalmia [40]. V. salmonicida causes Hitra disease in salmonids and cods and
contain marker protein VS-P1 on surface [41]. Many species have been reported to affect humans
also. V. alginolyticus reported for otitis media [42]; V. vulnificus for chronic hepatic diseases,
immunodeficiency symptoms, septicemia [43], purpuras, necrotizing dermo-hypodermatitis and
fasciitis [44] in patients; V. parahaemolyticus causes diarrhea, gastroenteritis and septicemia
[45,46]; and V. cholerae reported for cholera [47,48]. For diagnosis and identification many
molecular techniques (e.g. 16S rRNA, FISH, AFLP and RAPD) have been developed for these
species [49,50].
Flexibacteriosis
It is also named as Eroded mouth syndrome and Black patch necrosis and caused by
Tenacibaculum maritimum which has a wide host range, viz. turbot, gilthead sea bream, sea bass,
red sea bream, black sea bream sole, and salmonids [51]. Clinical symptoms are eroded and
hemorrhagic mouth, ulcerative skin lesions, frayed fins and tail rot [51]. T. maritimum is
antigenically and biochemically homogenous but two serotypes has been identified. Serological
identification, PCR based detection methods [52] and vaccination attempt have been made but an
effective vaccine is still awaited [53].
Pseudomonadiasis
Pseudomonas anguilliseptica caused mass mortality in eels of Japan, Taiwan, Scotland and
Denmark [54]. Its host range includes eels, black sea bream, ayu, wild herrings, gilthead sea bream
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and turbot. This pathogen causes red spot disease with clinical symptoms of abdominal distention
and hemorrhagic petechia in skin & lesions [55]. P. anguilliseptica has two ‘O’ serotypes, one of
eel and another of gilthead sea bream, turbot [55]. Clinical diagnostics and detection of the
pathogen by immunohistochemical and PCR methods is available [56,57]. Oil-adjuvant bacterins
were developed but any permanent inactivation is still awaited.
Streptococcosis
Cocci like Lactococcus garvieae, L. piscium, Streptococcus iniae, S. agalactiae, S. parauberis and
Vagococcus salmoninarum infect fishes throughout the world [58]. They infect species in Japan
(yellowtail, flounder); Spain, France, Italy, Israel (rainbow trout); USA (tilapia) and Australia
(barramundi). These cocci damage central nervous system with symptoms of exophthalmia and
meningo-encephalitis [59]. In humans Streptococcus iniae has been identified for cellulitis,
endocarditis, meningitis and septic arthritis [60]. Serotype studies revealed two serogroups (KGand KG+) in L. garvieae and two serotypes (I and II) in S. iniae [61]. S. agalactiae was reported
for illness and deaths in doctor fish (Garra rufa) [62]. Immunodiagnostic procedures are available
for identification of these cocci [63,64].
Diseases due to Flavobacteria
Flavobacteria are important aetiological pathogens represent by Flavobacterium psychrophilum,
F. branchiophilum and F. columnare, and cause bacterial coldwater disease (BCWD), bacterial
gill disease and columnaris disease respectively [65]. These species has wide host and geographic
ranges. CWD produce serious mortalities with clinical signs as increased mucus secretion,
epidermal hyperemia, splenomegaly and hepatomegaly [66,67]. Bacterial gill disease primarily
occurs in juvenile salmonids in spring season. Affected fish display lethargicity, high in water
column and gasping for air at surface. In columnaris disease pathological signs are erotic lesions
or deep ulcers on skin and fins. Proteome analysis of F. columnare has been performed that
exhibited virulence diversity in the species [68,69].
Edwardsiellosis
Two pathogenic species are in this group; Edwardsiella tarda and E. ictaluri. Among these E.
tarda has wide host range and reported to infect humans also [70] whereas E. ictaluri limited to
fishes only. These species cause systemic hemorrhagic septicemia and enteric septicemia in
catfishes [71]. A characteristic symptom observed is the presence of a raised or open ulcer on the
head. E. tarda causes loss of pigmentation, swelling on abdominal surface and petechial
hemorrhages in fins and skin [72,73]. Clinical diagnostic kits (i.e. ELISA, PCR and LAMP) are
available commercially to identify these species [74,75].
Enteric red mouth (ERM)
Yersinia ruckeri causes ERM which is responsible for critical economic losses. This bacterium
belongs to Enterobacteriaceae family and produce yersiniosis or ERM disease in fishes. Its
common host is Salmo gairdneri [76]. Clinical manifestations of this disease are exophthalmia,
ascites and hemorrhage & ulceration in jaw, palate, gills and operculum. Biochemically this
bacterium is very diverse and has two biotypes (1 & 2) grouped in many ‘O’ serogroups [77]. The
disease spread via direct contact with infected fish or by carriers. Progression of the pathogen in
infected rainbow trout has been demonstrated through bioluminescence imaging [78]. Recently
genetic component for holomycin resistance has been identified [79] and some probiotics and
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bacterins have also been developed for prevention of yersiniosis but permanent eradication still is
the matter of research [80,81].
Photobacteriosis
Also known as pasteurellosis, photobacteriosis is characterized by white nodules in spleen and
kidneys [82]. Photobacterium damselae subsp. piscicida is the causal agent. Mortality incidences
due to the photobacterium reported from Chesapeake Bay (white perch and striped bass), Japan
(yellowtail), Mediterranean countries of Europe (gilthead sea bream, sea bass and sole) and USA
[hybrid striped bass; 83]. Clinical diagnostics and laboratory identification (i.e. ELISA, PCR) are
available for photobacterium for disease control in farms [84].
Piscirickettsiosis
Piscirickettsia salmonis is responsible for piscirickettsiosis in rainbow trouts and salmons
principally in Coho salmon [85]. Observed clinical signs of this septicemic condition are nodules
in liver, darkening of the skin, lethargy, anorexia and respiratory distress in affected fish [86].
Commercial ELISA kits, PCR based protocols for diagnoses are available in the market [87].
Winter ulcer
It’s a winter season problem characterized by skin ulcers at scale covered surface; mainly occur in
Atlantic salmons with few incidences reported in plaice and rainbow trouts also [88]. Causal
pathogen for winter ulcer is Moretella viscosa which is represented by two groups: first one is
Norway group and second one is Iceland group [89,90].
Table 1: Bacterial pathogens in fishes.
Bacterium
Genome
size
(Mb)
4.12
Disease
Main host(s)
Vaccine(s) if any
Vibriosis
V. ordalii [92]
3.41
Vibriosis
Salmonids, cods,
red sea bream
Salmonids
V. vulnificus [93]
Aliivibrio salmonicida
[41]
Flavobacterium
psychrophilum [67]
F. branchiophilum [94]
F. columnare [95]
5.26
4.65
Vibriosis
Vibriosis
Eel, tilapia
Atlantic salmon
2.86
Coldwater disease
Salmonids, carp
Vibrio anguillarum-ordaliiYersinia ruckeri Bacterin
Vibrio anguillarum-ordaliiYersinia ruckeri Bacterin
-----Aliivibrio salmonicida
Bacterin
-------
3.56
3.2
Bacterial gill disease
Columnaris disease
Edwardsiella tarda [96]
3.76
Edwardsiellosis
Edwardsiella ictaluri
[97]
Aeromonas hydrophila
[98]
A. salmonicida [18]
A. veronii [99]
3.81
Enteric septicemia
Reared fishes
Cyprinids,
salmonids
Salmons, catfish,
carps
Catfish
4.74
Renibacterium
salmoninarum [25]
3.15
Motile aeromonas
septicemia
Furunculosis
Epizootic ulcerative
syndrome
Bacterial kidney disease
Salmonids, non
salmonids
Salmons
Salmonids, non
salmonids
Salmonids
Vibrio anguillarum [91]
4.70
4.55
------------------Edwardsiella ictaluri
Bacterin
Aeromonas hydrophila
Bacterin
A. salmonicida Bacterin
---------------
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Streptococcus
parauberis [100]
Lactococcus garvieae
[101]
Mycobacterium
marinum [102]
Yersinia ruckeri [103]
2.14
Streptococcosis
Turbot
--------
2.17
Lactococcosis
Yellowtail, eel
--------
6.64
Mycobacteriosis
--------
3.58
Enteric redmouth
Photobacterium
damselae [104]
5.05
Photobacteriosis
Sea bass, turbot,
Atlantic salmon
Salmonids, eel,
minnows
Sea bream, sea
bass, yellowtail
Yersinia ruckeri Bacterin
--------
Fungal diseases in fish
Fungal diseases are quite rare than bacterial and viral ones in mass mortality means. Fungi can
infect fishes if they are given environmental stress conditions like poor environmental conditions
(water with low circulation, low dissolved oxygen or high ammonia or high organic loads), poor
nutrition or injury. In most cases fungal infection occurs as secondary infection in association with
bacterial or viral. Common fungal infections are due to water molds (Saprolegnia). This cotton
wool fungus belongs to Saprolegniaceae family and causes saprolegniasis in fishes and fish eggs
[105]. Saprolegnia grow optimum at temperature ranges 59o to 86o F [106]. In most cases
saprolegnia attacks on existed injured tissue and spread to healthy tissues [107]. These saprophytes
transmit via motile zoospores and detritus of bottom. Saprolegniasis symptoms are superficial and
observed as fluffy tufts of cottony growth on the skin and gills in affected fish [108].
Another common fungal disease in fishes is gill rot (Branchiomycosis) that occurs mainly in carp
and eels. Its causative agents are Branchiomyces sanguinis and B. demigrans. These species
invades fish in water of low pH, low dissolved oxygen and high algal bloom. Infected fish manifest
respiratory distress, thrombosis in blood vessels of gills and necrosis in gills [109]. Another fungal
problem in fish is Icthyophonus disease caused by Icthyophonus hoferi [110]. Icthyophonus
manifest itself internally and primarily attack liver and kidney [111]. Disease is transmitted via
fungal cysts released by faeces and via cannibalism of infected fish [112]. A fish fungus is
Exophiala [113] which is represented by two important pathogenic species namely E. salmonis and
E. psychrophila [114]. These fungi produce round yellow to white granulomas in liver, kidneys
and spleen with enlargement of posterior kidney [115]. To manage fungal diseases sanitation of
aquatic system, avoidance of overcrowding to minimize injury, use of anti-fungal agents and good
nutrition are things to keep in mind.
Viral diseases in fish
Infectious hematopoietic necrosis (IHN)
Aetiological agent of IHN is infectious hematopoietic necrosis virus (IHNV). Its principle host is
rainbow trout (Oncorhynchus mykiss) and this virus causes IHN in trouts and salmonids. IHNV,
endemic in salmons of North America, emerged as pathogen in farmed rainbow trouts in USA in
1970 [116]. Thereafter, it expanded its geographical and host range by increasing virulence and
pathogenicity [117] and spread worldwide via contaminated eggs. It is an enveloped bullet shaped
virus that belongs to genus Novirhabdovirus with in Rhabdoviridae family [118] and contain –ve
sense ssRNA of approximate 11000 nucleotide length [119]. Its genome contain six genes (N-, PInternational Journal of Microbiology and Allied Sciences, Nov 2014, Volume 1 Issue 2
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, M-, G-, NV- and L-genes) located from 3’ to 5’ [120]. IHNV infects many economically
important species of aquaculture; principally of genus Oncorhynchus, several salmonids and nonsalmonid species [121]. This virus proliferates in vascular endothelial tissues, nephron cells and
hematopoietic tissues which results in viraemia, impairment of osmotic balance, edema and
hemorrhages. IHNV produced severe aquaculture losses in USA, Canada, Japan, Russia, Austria,
Czech Republic, France, Germany, Italy, Netherlands, Poland and Spain through infected eggs
and/ or fish [122]. Phylogenetic studies identified 5 genogroups (U, M, L, E and J) and multiple
subtypes exist in each genogroup that produce diversity in virulence and pathogenicity of virus
[123]. This virulence diversity has been demonstrated in rainbow trout [117]. In North America U,
M and L genogroups occurs [124]. Molecular epidemiology and evolution of IHNV has been
assessed using Bayesian analyses of N, G and NV genes [125]. Diagnostic and identification
methods have been developed and validated using reverse transcriptase PCR of nucleocapsid gene
[126]. Transmission of virus takes place horizontally, vertically, by biological vectors such as fish
parasites (e.g. Piscicola salmositica, Salmonicola sp.) and by carrier fish [127]. IHNV is released
in faeces, urine and external mucus. In persistently infected rainbow trout, truncated virion
particles were detected; those were proved to be mediators of persistence [128]. Several vaccine
trials have been perfomed [129], as IHNV rapidly alter its virulence by evolution under selection
pressure [122,130].
Viral hemorrhagic septicemia (VHS)
Causative agent is viral hemorrhagic septicemia virus (VHSV) which was initially assumed to be
endemic in Europe in rainbow trout but further studies indicated the larger host and geographical
range of this virus [131]. It is an enveloped ssRNA virus belonging to Novirhabdovirus genus of
Rhabdoviridae family [132]. Its genome (11 kb) has been sequenced [133] and comprises six open
reading frames (3’-N-P-M-G-NV-L-5’) encoding N-, P-, M-, G-, NV- and L- proteins [134]. Many
finfish species are susceptible to VHSV like species of family Clupidae, Pleuronectidae,
Salmonidae, Esocidae and Gadidae [15]. Multiplication of the virus takes place in endothelial cells
of blood capillaries (leading to hemorrhagic lesions in internal organs and musculature),
leucocytes, hematopoietic tissues and nephron cells. Clinical symptoms in affected fish are
hemorrhages in muscles and liver. Externally, fish may show hemorrhage on skin, around eye orbit
or exophthalmia in eye. Cytopathic effects of VHSV have been assessed on macrophage cell line
[135]. VHSV is very diverse in virulence & pathogenicity [136]. Till now four of strains (I-IV)
have been identified in VHSV with several sub strains [137] and most strains limited to northern
hemisphere. Strain I comprises European isolates; strain II include isolates from the Baltic Sea;
strain III consists of North Atlantic Ocean isolates; strain IV involve Japanese/ Korean (IVa) and
North American (IVb) isolates. In 1998, thousands of pacific herring, hakes and walley pollock
were found dead in Alaska due to American strain of VHS [138]. Pathogenicity in pacific herrings
and Atlantic salmon [139] and phylogenetic diversity & biogeography of VHSV has been assessed
using Bayesian approach [140]. A few rapid and accurate methods to detect VHSV are available
[141]. In developed countries like UK, US, Norway, Denmark; effective control has been achieved
but due to dynamic pathogenicity of the virus permanent eradication is still awaited [142].
Spring viraemia of carp (SVC)
Spring viraemia carp virus (SVCV) causes SVC in carps (Cyprinus carpio). Outbreaks usually
occur in the spring time, when water temperature ranges between 10-17 oC [143]. SVCV belongs
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to genus Vesiculovirus and it lacks the nonvirion gene [144]. Its genome consists of one molecule
of ssRNA that has been completely sequenced [145] and molecular analysis & virulence diversity
have been addressed of the virus [146]. Virion particle has an inner nucleocapsid that is surrounded
by a lipid containing envelope with spikes. Viral RNA encodes five structural proteins; L-, G-, P, N- and M- proteins [145]. SVCV isolates are classified in 4 genogroups; Ia, Ib, Ic and Id [147].
Strain Ia occurs in US and England, Ib in Ukraine [148] and Moldova [149], strain Ic in Russia
[150] and Id in UK. SVCV is mainly associated with losses in common carp and koi carp in spring
season and can infect many fish species [15]. Important disease symptoms include exophthalmia,
abdominal swelling, hemorrhages in the skin, gills & eyes and darkening of the skin [150].
Pathogenesis of SVCV infection and related proteins expression was assessed on epithelioma
papulosum cyprini cells [151]. For diagnostics; antibody detection and PCR methods are available
for epidemic alert [152]. SVCV shed in faeces, urine and sexual fluids and transmit via contact and
animate vectors such as Argulus foliaceus, Piscicola geometra [144]. Attempts for DNA vaccines
were made and permanent vaccine development is however still in progress [153].
Infectious salmon anemia (ISA)
Infectious salmon anemia virus (ISAV) is the infectious agent for ISA in salmons (Salmo salar).
ISA was first recognized in Atlantic salmons from Norway [154]. In 1996, ISA cases were reported
from Canada [155]. There ISAV was found to be associated with hemorrhagic kidney syndrome
(HKS) in Atlantic salmon [156]. Thereafter in 1998, ISA was detected in Scotland [157].
Subsequent reports came from USA [158], Chile [159], and Faroe Islands [160]. The virus was
then isolated from salmons and rainbow trout on SHK-1 cell line and well characterized [161].
ISAV belongs to the family Orthomyxoviridae and genus Isavirus [162]. It is an enveloped
pleomorphic virus with eight segments of –ve sense ssRNA as genome (14.5 kb) which encodes
hemagglutinin-esterase (HE) [163], a surface glycoprotein, responsible for strain virulence [164].
ISAV produce serious losses in aquaculture especially in salmons [161]. Systemic infection is
characterized by anemia, ascites, congestion and enlargement of the liver and spleen [159]. ISAV
is divided in two groups on the basis of HE gene; the European group and North American group
[164]. Molecular studies revealed high polymorphism regions (HPR) in the virus [162].
Pathogenesis and cell tropism in atlantic salmon gill epithelial cells has been demonstrated on
SHK-1, ASK-II and TO cells using IFAT [165]. Clinical diagnostic and laboratory identification
methods (in situ hybridization, RT-PCR, IFAT, cell lines) are available for virus [166]. The virus
transmit horizontally, vertically, by sea lice (Caligus elongates, Lepeophtheirus salmonis) or via
coprophagy [167]. For disease control several vaccination field trials were performed and effective
vaccines are available in sensitive zones; North America, Faroe Islands and Norway [168].
Alphavirus-based replicon expression vectors are gaining very well acceptance for recombinant
vaccine development [169]. HE gene has been confirmed as potential candidate gene for
vaccination and a HE based alphavirus-replicon expression system has been developed against
ISAV [170].
Epizootic hematopoietic necrosis (EHN)
Epizootic hematopoietic necrosis virus (EHNV) is the causal agent of this necrosis and was initially
identified in red fin perch (Perca fluviatilis) and rainbow trouts in Australia [171]. EHNV is a
member of Iridoviridae family and placed in the genus Ranavirus [172]. Virion particles are
icosahedral and retain dsDNA as genome [173]. It mostly infects red fin perch [174]. Some other
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species like Esox lucius, Sander lucioperca and Ameiurus melus are also susceptible to EHNV
infection [175]. Susceptibility of some other native and non-native species of southeast Australia
towards EHNV has also been confirmed [176]. EHNV causes severe necrosis in hematopoietic
tissues in perch and rainbow trout and create great loss in aquaculture production. Histopathology
of infection involves necrosis in liver, spleen & renal hematopoietic tissues [177] and ulcerative
dermatitis and swim bladder edema are observed additionally in rainbow trout [172].
Immunodiagnostic and PCR methods have been developed and validated for EHNV [178,179].
EHNV spread via spill over of virus from endemic reservoir [172]. To date no effective biosecurity
measures are available for EHN control, a permanent vaccine development against EHN is still
awaited.
Mortalities due to Red sea bream iridovirus (RSIV)
RSIV was first identified in red sea bream (Pagrus major) of Japan [180] and thereafter mortality
outbreaks were reported worldwide [181]. It was isolated from kidneys and spleen of infected fish
using cell cultures and well characterized [182]. It is a member of genus Megalocytivirus within
Iridoviridae family [183]. Other megalocytiviruses were also reported time to time in severe
outbreaks with necrosis; infectious spleen and kidney necrosis iridovirus in mandarin fish [184],
sea bass iridovirus, rockbream iridovirus [185] and orange spotted grouper iridovirus [186].
Complete genome of RSIV has been sequenced and phylogenetic analysis was performed using
major capsid protein and ATPase genes [187]. This virus affects especially juveniles of red sea
bream. Infected fish manifest anemia, petechia of the gills and enlargement of the spleen [188].
Pathogenicity of the RSIV was demonstrated in Japanese amberjack (Seriola quinqueradiata)
[189]. Histopathology of affected tissues showed development of inclusion body bearing cells that
are specific characteristic of RSIV infection [190]. Antigenic and molecular studies revealed three
lineages in megalocytiviruses [172]. Clinical diagnostics and detection strategies (e.g. serological
analysis, RT-PCR, cell culture, immuno-fluorescence) have been established for RSIV [191].
Experimental efforts for DNA vaccines were attempted time to time and a formalin inactivated
vaccine is commercialized against RSIV [192]. A recombinant vaccine (351R-GADPH fusion
protein) has found to be an effective immunogenic intervention for RSIV [193].
Viral nervous necrosis (VNN)
Nervous necrosis virus (NNV) causes severe nervous necrosis in wide range of economically
important species [194]. Although initially described in Australia in barramundi [195], this disease
was subsequently identified in turbot in Norway [196], sea bass in Martinique [197] and parrotfish
& red spotted grouper in Japan [198]. Thereafter the virus was isolated by cell culture on SSN-1
cell line [199] and genetic studies well characterized as member of genus Betanodavirus within
Nodaviridae family [200]. Virion particles are spherical, non-enveloped and retain two +ve sense
ssRNA (RNA1 and RNA2) molecules [201]; among which non-structural proteins are the product
of RNA1 and coat proteins are encoded by RNA2 [202]. VNN create serious economic loss in
aquaculture worldwide especially in larvae and juveniles [203]. This RNA virus causes
encephalopathy and retinopathy in marine fishes such as sea bass, halibut, striped jack and grouper
[204]. Nervous necrosis is characterized by vacoulation of neural cells of brain, retina and spinal
cord [205] due to which affected fish lost its balance and reflects muscular and visual dysfunction.
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Histopathology and virulence diversity of NNV has been demonstrated in larvae of humpback
grouper (Cromileptes altivelis) [206]. Betanodaviruses affect wide range of host species [207] and
to date reported from south and Mediterranean Europe and Tunisia, Norway, East Asia, Oceania,
UK, and North America. Molecular phylogenetics revealed four clades in the virus as striped jack
NNV, tiger puffer NNV, bar fin flounder NNV and red grouper NNV [208]. Phylogenetic studies
have also been performed in larva of clownfish Amphiprion seabae Bleeker [209]. Genome
sequencing coupled with transcriptome analysis (using microarray) reveal some candidate immune
genes during viral infection for vaccine development [210]. Transcriptome profiling of viral
infection has also been recently demonstrated in Atlantic cod brain [211]. Rapid and convenient
RT-PCR methods have been developed for diagnosis and epidemic alert [212]. NNV infect actually
larval stage, in which vaccination is quite difficult however potentiality of some peptide vaccine
(virus like particles; VLP) and live vaccines have been investigated [213,214]. VLPs are proved to
be effective vaccine candidates as they elicit efficient antibody response [215]. A recombinant
RGNNV capsid VLP has been recently purified [216].
Infectious pancreatic necrosis (IPN)
IPN was first recognized in brook trout (Salvelinus fontinalis) at West Virginia, USA [217] and
was the first fish virus to be isolated via cell culture approach [218]. In the following years IPNV
strains were recognized in many countries around the world [219]. This virus can transmit by
faeces, urine and sexual products of infected fish and it easily spread by shipment of contaminated
fish eggs from one country to another [220]. It may also spread by faeces of piscivorous birds, viz,
heron, crow, grackle, kingfisher, sparrows, mallards, egrets and ospreys [221]. Now IPNV is
probably present in all major trout farming countries and mainly infect fry and juveniles of rainbow
trout, brook trout and atlantic salmon. It is an icosahedral, non enveloped aquabirnavirus [217] and
very diverse in host range affecting fishes from freshwater to mariculture [222]. To date 4
serogroups (A, B, C, D), and 9 serotypes (A1-A9) has been identified within serogroup A which
are antigenically very diverse [223]. A1 serotype exists in USA; A2, A3, A4, A5 & A10 in Europe;
and A6-A9 are reported from Canada. Serotypes A1, A2 and A3 have been reported from Asia
also [224]. Virion particles retain two segments (A and B) of dsRNA which encode 5 viral proteins.
Segment B encodes VP1 protein (RNA polymerase) and segment A encodes VP2, VP3, VP4 and
VP5 proteins. Among these VP2 is expressed as epitope that determine serotype specificity [225].
VP3 is an inner structural protein that bound to RNA and create ribonucleoprotein core of the virus
[226]. VP5 protein regulates apoptosis programming of host cell. Virulence diversity (in VP2)
especially serotyping and host range among them hindered the progress of control & biosecurity
measures [227]. Recently stress induced virulence dynamics in IPNV has been studied in atlantic
salmon naive fry [228]. IPNV has been demonstrated to induce apoptosis preceding necrosis in
fish cell lines by down regulating the survival factor Mcl-1, a Bcl-2 homologue [229]. The virus
can be identified by laboratory procedures like RT-PCR, flow cytometry, cell culture and
histopathology [230]; and immunohistochemistry of viral infection has been demonstrated in
cultured rainbow trout [231]. IPNV reside in leucocytes and reduce immunity component secretion
from host cell. Innate immune components after viral infection are well described in salmonid fish
[232]. Several kinds of prevention & disease control methods are available for the pathogen and
vaccines have been commercialized against the virus [233]. Immunogenicity and antibody
components evoke against viral infection in Atlantic salmon, has been described in detail by
Munangandu and coworkers [234,235].
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Losses due to salmon alphaviruses
Some alphaviruses also pose serious threats to aquaculture [236]. Alphaviruses belongs to
Togaviridae family and cause sleeping disease and pancreas disease in rainbow trouts and Atlantic
salmon respectively. Salmon alpha virus (SAV) was identified as aetiological agent for outbreaks
in Ireland [237], Norway [238] and Scotland [239]. Sequence analyses of SAV revealed six
subtypes (SAV1-6) in the pathogen based on E2 and nsP3 genes [240]. These subtypes are reported
from Europe [241], Norway [242], Ireland, England, Germany, France, Italy, Spain and Scotland
[240]. Polyprotein sequences of Scottish and Irish isolate was compared recently in which Irish
isolate showed high viral load and cytopathic effects in cell culture [243]. Clinical signs of this
disease are cardiac myopathy and degeneration of exocrine pancreas [244]. Alphaviruses transmit
by arthropods and insects (principally mosquitoes) [245]. Virion particles are spherical, enveloped
and retain 12 kb long +ve sense ssRNA [246]. Full genome has been sequenced of some subtypes
[247]. Recently some Norwegian SAV3 strain’s genomes have been sequenced that revealed
defective viruses with some genome deletions [248]. Host pathogen interactions and innate &
adaptive immune responses against SAV-3 Norwegian strain has been well characterized in
Atlantic salmon [249]. Subunit vaccines (of spike proteins E1, E2), DNA vaccine and inactivated
whole virus vaccines have been developed for SAV control [250]. And, inactivated whole virus
vaccine has been proved better immunogenic when compared to spike protein vaccine or DNA
vaccine [251].
Diseases due to herpesviruses
Herpesviruses also produce a great loss in aquaculture production. Important members include
oncorhynchus masou virus (OMV), ictalurid herpes virus (IcHV) and koi herpes virus. OMV is
also known as salmonid herpesvirus (SaHV) and belongs to the Herpeseviridae family. This family
includes enveloped quasispherical viruses and retains a linear dsDNA as genome. OMV was
isolated in Japan from ovarian fluid of masou salmon (Oncorhynchus masou). Virus multiplies in
endothelial cells of blood capillaries, hematopoietic tissues and hepatocytes; and produce
epithelioma around upper and lower jaw. Intestinal hemorrhage and white spot on liver are
additional internal signs. It is reported to be an oncogenic virus for fishes and produce skin
ulcerative condition in affected salmon. The DNA of SaHV was shown to molecular weight of
about 100×106 Da [252]. SaHV shed by horizontal and vertical transmission [253]. In the survivors
of acute infection, epidermal papillomas are developed [254]. This virus can be identified by direct
methods as neutralization tests IFAT or ELISA [15]. Formalin-inactivated OMV vaccine is
recommended for prevention [255].
KHV belongs to Cyprinivirus genus within family Alloherpesviridae [256] and is responsible for
explosive losses in common carp and koi carp [257]. It was first recognized in UK [258] and then
reported worldwide in carp industry [259]. Recently, KHV has been detected in South Korea also
[260]. KHV is an enveloped dsDNA virus [261] with genome size 295 kb [259] and persists
initially in latent or carrier phase in infected fish without exhibiting any clinical signs [262].
Clinical symptoms may include gill mottling with red and white patches, sunken eye, pale patches
on skin and notched nose [263]. Affected fish may exhibit hyper secretion of mucus in severe stage
of infection. Detection methods for KHV like PCR, ELISA are available for diagnosis [264]. A
safe and effective vaccine is currently not available for KHV. Recently, some anti-KHV bacteria
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have been isolated from carp intestine and their habitat water [265]. A DNA vaccine comprising
ORF25 fragment has been constructed and evaluated for immunogenicity [266].
IcHV is also known as channel catfish herpesvirus (CCHV). It is responsible for economic losses
of channel catfish (Ictalurus punctatus), blue catfish (Ictalurus furcatus) and white catfish
(Ictalurus catus) fry and fingerlings in North America, Russia and Honduras [267]. IcHV genome
is 137.2 kbp long and has 77 ORFs [268]. The envelope glycoprotein is encoded by ORF 59 [269].
From all of the ORFs, ORF 50 encodes a secreted, mucin like glycoproteins (gp250) due to which
there are different isolates of IcHV [270]. A thymidine kinase (TK) is encoded by ORF 5 [271]. It
was demonstrated that IcHV virulence is reduced at temperature below 20 oC. Diagnostic and
identification can be done by IFAT, neutralization test, ELISA [272] or by PCR [273].
Recombinant TK gene deletion mutants of IcHV were constructed and exposure of TK- mutants to
fish induces protective immunity against challenge [274].
Table 2: Viruses in shrimps and their geographical distribution
Virus
Taura syndrome virus (TSV)
Genome
(+) ssRNA
Geographical distribution
Asia, Americas
(+) ssRNA
(+) ssRNA
Family
Dicistrovirida
e
Roniviridae
Nodaviridae
Yellow head virus (YHV)
Macrobrachium rosenbergii nodavirus
(MrNV)
Infectious myonecrosis virus (IMNV)
Infectious hypodermal and
hematpoietic necrosis virus (IHHNV)
Hepatopancreatic parvovirus (HPV)
White spot syndrome virus (WSSV)
Baculoviruses
(+) ssRNA
ssDNA
Totiviridae
Parvoviridae
Indonesia, china, brazil, Thailand
Asia, Africa, Americas
ssDNA
dsDNA
dsDNA
Parvoviridae
Nimaviridae
Baculoviridae
Asia, America, Africa, Madagascar
Asia, America
Asia, Africa, America, Australia
Asia, Mexico
Asia, Australia
Global impacts of fish diseases and pathogens
Emerging diseases in aquaculture have crucial effect on country economy especially of coastal
nations (Table 3). Like in 1996, 40% of total shrimp production in US was affected due to viral
diseases that were estimated up to US $ 3 billion [275]. Major significant production loss was
always immediate after any pathogen has emerged and has pronounced impact till on fish/ shrimp
industry till any biosecurity measure developed. Poor, uneducated, low income rural farmers are
critically affected. WSSV has been estimated the worst shrimp pathogen as produced US $ 4-6
billion loss in Asia till ten years after its emergence [276]. The loss to fish based industry because
of diseases caused by IHHNV and TSV is estimated to be approximately US $ 2 billion [277].
ISAV produce 30 million dollars loss in Scotland and approximate 20 million in Norway and
Canada within one year after its emergence. In 2010, ISA hit Chilean salmon industry by
decreasing their production from 600,000 (in 2008) to 100,000 metric tonnes [278]. Another
example of ISA loss was of British Columbia in 2011 [279]. KHV produced critical loss in
Indonesia [280]. These pathogens account for severe impact on susceptible species and their
biodiversity by increasing host range, changing genetic diversity and affecting predator/prey
animals [281]. For farming, aquatic animals are transferred actually to stress environment in which
they are exposed to artificial feeding, poor water quality and high stocking density. These factors
facilitate pathogen to compromise host defense system, replicate in the susceptible hosts and thus
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contribute to disease transmission [282]. Expanding world trade of aquatic animals contributes to
increase in geographical range for pathogens [283].
Table 3: Geographic prevalence of notifiable viruses in fishes and vaccines developed against
them [15].
Virus
Vaccine(s) and manufacturer(s)
VHSV
IHNV
----------Aqua Health Ltd. Novartis Canada
SVCV
Bioveta Czech Republic
EHNV
ISAV
----------FORTE VI Aqua Health
Novartis Canada;
Ltd.
Targeted antigen(s)
----------G protein
Geographic distribution
Europe, North America, Asia
Europe, North America, Asia
Inactivated virus
Americas, Europe, Asia
----------Inactivated virus
Australia
Norway, Canada, USA, Chile,
UK, Faroe islands
North America, Europe, Asia,
Australia, Africa
Asia
Americas, Europe, Asia
Microtek International Inc. British
Columbia Canada
NNV
-----------
-----------
RSIV
IPNV
----------AquaVac IPN Oral Intervet
International BV Netherland
----------VP2 and VP3
KHV
-----------
-----------
SAV
PD, Pharmaq AS Norway;
Inactivated virus
Norvax Intervet International BV
Netherland
OMV
------------
-----------
Asia, Africa, Europe, north
America, Israel
Europe, Norway, Ireland,
England, Germany, France,
Italy, Spain, Scotland
Japan
Diseases in fish or shrimps are caused by natural infection or spill-over from carrier reservoir [284].
Pathogens of these diseases especially the viral ones, have emerged in new geographical region
due to their global trade [285]. IHHNV spreads throughout Americas by translocation of infected
P. monodon from Philippines [286]. ISAV emerged in farmed Atlantic salmons worldwide from
wild species of Norway and Canada [287]. IHNV, VHSV, SVCV and KHV spread Europe, Asia
and Americas as result of prolific international trade. Similarly, there is wider transmission of many
viruses like WSSV, TSV and IMNV which as devastating pathogens for shrimps [276,288]. So the
present scenario demands effective vaccines against the mentioned pathogens for sustained
aquaculture. To date vaccines have been developed for some economic bacterial & viral pathogens
only [289] and no permanent biosecurity procedure is available for fungal & parasitic diseases.
Vaccines can be typed as inactivated, attenuated and recombinant subunit & DNA vaccines. An
ideal vaccine must be multivalent, orally deliverable and must provoke strong long lasting immune
response. Recombinant subunit & DNA vaccines are potential in this regard. Most of the vaccines
developed yet are delivered by intraperitoneal injection but it is an expensive, labor intensive and
time consuming process. So efforts are in progress to develop orally deliverable vaccines via
feeding route. Factors like increasing demand for seafood, global population and limited
production from fisheries enhancing the global trade of aquaculture to complete this demand; that
eventually led to emergence of above mentioned and new diseases in finfish & shrimp aquaculture.
Development of cell lines resulted in enhanced surveillance of aquatic virosphere. Recombinant
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technologies and analytic advanced techniques may play a promising role to overcome aquaculture
economic losses. Genome sequences of the fish pathogens provide us information about protein
expression in host which may represent key virulent proteins as suitable target(s) for vaccine
development. Understanding of molecular and metabolic mechanisms will indeed improve our
knowledge in vaccine development against these pathogens.
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