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Viral Encephalopathy and Retinopathy in Aquaculture: a Review Q.K CORE Metadata, citation and similar papers at core.ac.uk Provided by Archive Ouverte en Sciences de l'Information et de la Communication Viral encephalopathy and retinopathy in aquaculture: a review Q.K. Doan, Marc Vandeputte, B. Chatain, T. Morin, F. Allal To cite this version: Q.K. Doan, Marc Vandeputte, B. Chatain, T. Morin, F. Allal. Viral encephalopathy and retinopathy in aquaculture: a review. Journal of Fish Diseases, Wiley, 2017, 40 (5), pp.717-742. 10.1111/jfd.12541. hal-01533903 HAL Id: hal-01533903 https://hal.archives-ouvertes.fr/hal-01533903 Submitted on 26 May 2020 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Journal of Fish Diseases 2017, 40, 717–742 doi:10.1111/jfd.12541 Review Viral encephalopathy and retinopathy in aquaculture: a review Q K Doan1,2, M Vandeputte1,3, B Chatain1, T Morin4 and F Allal1 1 Ifremer, UMR 9190 MARBEC, Palavas-les-Flots, France 2 TNU, Thai Nguyen University of Agriculture and Forestry (TUAF), Quyet Thang Commune, Thai Nguyen City, Vietnam 3 INRA, GABI, AgroParisTech, Universite Paris-Saclay, Jouy-en-Josas, France 4 Anses, Ploufragan-Plouzane Laboratory, Unit Viral Diseases of Fish, Plouzane, France expected to induce the spread of pathogens towards Abstract higher latitudes and to provoke negative impacts on Viral encephalopathy and retinopathy (VER), fish physiology (Cochrane et al. 2009). Among otherwise known as viral nervous necrosis (VNN), others, the viral encephalopathy and retinopathy is a major devastating threat for aquatic animals. (VER), otherwise known as viral nervous necrosis Betanodaviruses have been isolated in at least 70 (VNN), is considered one of the most serious viral aquatic animal species in marine and in freshwater threats for almost all marine aquaculture fish species environments throughout the world, with the and requires a special focus due to the fact that out- notable exception of South America. In this breaks mostly happen in warm conditions. This dis- review, the main features of betanodavirus, includ- ease, detected in at least 70 cultured or wild marine ing its diversity, its distribution and its transmis- and fresh water species, already caused serious eco- sion modes in fish, are firstly presented. Then, the nomic losses in the aquaculture industry in the past existing diagnosis and detection methods, as well decades, and we can anticipate larger impacts of this as the different control procedures of this disease, disease because of global warming. are reviewed. Finally, the potential of selective No simple and effective procedures are available breeding, including both conventional and geno- to treat this disease in fish. It is, therefore, impor- mic selection, as an opportunity to obtain resistant tant to develop tools and set up new approaches commercial populations, is examined. to limit the occurrence and impacts of VNN epi- sodes in aquaculture farms. Keywords: betanodavirus, disease resistance, genetics, To stress that need, we present here an exten- nervous necrosis virus, selective breeding. sive review about VNN disease in aquaculture, including the features of the virus, the available procedures to control this disease and the poten- Introduction tial of selective breeding and genomic selection (GS) for resistance to viral diseases, as a prospec- Although there is presently no strong evidence high- tive way to prevent VNN disease in fish. lighting a possible raise of fish disease outbreaks due to climate change, increasing temperatures are Nervous necrosis virus Correspondence Q K Doan, Ifremer, UMR 9190 MARBEC, Chemin de Maguelone, 34250 Palavas-les-Flots, France The causative agent of VNN, the nervous necrosis (e-mail: [email protected]) virus (NNV), was classified as a member of the Ó 2016 John Wiley & Sons Ltd 717 Journal of Fish Diseases 2017, 40, 717–742 Q K Doan et al. Viral nervous necrosis in aquaculture Nodaviridae family (Mori et al. 1992), which con- recorded to be infected by betanodaviruses (Mun- tains two genera: alphanodavirus and betano- day et al. 2002; Shetty et al. 2012). davirus (Van Regenmortel et al. 2000). The species of the first genus were originally isolated Molecular structure from insects (Fig. 1), but appear to infect both vertebrates and invertebrates, and to cause the Betanodavirus contains a bisegmented genome death of insect and mammalian hosts (Adachi composed of two single-stranded, positive-sense et al. 2008). Betanodaviruses usually affect the RNA molecules (Mori et al. 1992). The nervous system of marine fish, leading to beha- sequence of RNA1 is about 3.1 kb and includes vioural abnormalities and extreme high mortalities an open reading frame (ORF) encoding a RNA- (Munday, Kwang & Moody 2002). In mammals, dependent RNA polymerase (RdRp) of 110 kDa the pathogenicity of betanodaviruses is poorly catalysing the replication of the virus, also reported, but mice have been demonstrated as named protein A (Nagai & Nishizawa 1999). non-susceptible, and human cells as not permeable The sequence of RNA2 (1.4 kb) encodes the to that genus (Adachi et al. 2008). Recently, a capsid protein (37 kDa), which may have a new emerging disease, the white tail disease function in the induction of cell death (Guo (WTD), which affects the giant freshwater prawn et al. 2003). In addition, during the virus repli- and the whiteleg shrimp Penaeus vannamei, has cation, a subgenomic RNA (RNA3) is synthe- been demonstrated to be caused by the Macro- sized from the 30-terminus of RNA1 (Ball & brachium rosenbergii nodavirus (MrNV). Sequence Johnson 1999). This RNA3 encodes two other analysis of this virus suggests the existence of a non-structural proteins, B1 (111 amino acids) new genus, gammanodavirus, infecting crustaceans and B2 (75 amino acids). Protein B1 displays (Qian et al. 2003; Senapin et al. 2012, Fig. 1). antinecrotic property enhancing the viability of viral host cell (Sommerset & Nerland 2004). Protein B2 is an inhibitor of host RNA silencing General morphology in either alphanodavirus or betanodavirus, but Betanodavirus virions were first described as non- could also promote mitochondrial fragmentation enveloped, spherical in shape and have icosahedral and cell death induced by hydrogen peroxide symmetry, with a diameter around 25 nm and a production (Su et al. 2014). capsid formed by 180 copies of a single protein of 42 Kda (Mori et al. 1992). A similar virus of 20– Classification 34 nm in diameter was detected in infected Asian sea bass Lates calcarifer larvae, striped jack Pseudo- Betanodavirus was described for the first time caranx dentex, turbot Scophthalmus maximus, from infected larval stripped jack. The name European sea bass Dicentrarchus labrax (Glaze- striped jack nervous necrosis virus (SJNNV) was brook, Heasman & Beer 1990; Yoshikoshi & consequently adopted (Mori et al. 1992). Subse- Inoue 1990; Bloch, Gravningen & Larsen 1991; quently other agents of VNN were isolated from Munday et al. 1992), and many various fish spe- diseased fish species (Munday et al. 2002). The cies throughout the world were subsequently first comparative studies between viral strains iso- lated from different marine fish species were car- ried out in the middle of the 1990s, where Nishizawa et al. reported the sequence of SJNNV and four different fish nodaviruses as well as four different insect nodaviruses (Nishizawa et al. 1995). From a phylogenetic analysis of the RNA2 T4 variable region, betanodaviruses were classified into four different species designed as the SJNNV type, the barfin flounder nervous necrosis virus (BFNNV) type, the red-spotted grouper nervous necrosis virus (RGNNV) type and the tiger puffer nervous necrosis virus (TPNNV) type (Nishizawa Figure 1 Three genera of Nodaviridae. et al. 1997). These species partially correlate with Ó 2016 John Wiley & Sons Ltd 718 Journal of Fish Diseases 2017, 40, 717–742 Q K Doan et al. Viral nervous necrosis in aquaculture three different serotypes determined from virus An alternative classification has been proposed neutralization using polyclonal antibodies (sero- (Thiery et al. 2004). However, this numerical type A for SJNNV species, B for TPNNV species nomenclature (cluster I, II, III and IV), indepen- and C for BFNNV and RGNNV species) (Mori dent from the host species origin, is not exten- et al. 2003). Each species corresponds to different sively used because viruses from different clusters host fish and different in vitro optimal growth could infect a same host species, for example temperatures (Table 1). RGNNV is the most European sea bass (Thiery, Raymond & Castric popular species because a variety of fish species, 1999a; Thiery, Arnauld & Delsert 1999b), and distributed in warm water, are affected (optimal the classification was not consistent with geo- growth temperature of 25–30 °C) (Asian sea bass, graphical areas (Dalla Valle et al. 2001; Thiery European sea bass, groupers), whereas BFNNV is et al. 2004; Cutrın et al. 2007). restricted to cold-water (15–20 °C) marine fish species (Atlantic halibut Hippoglossus hippoglossus, Phylogenetic relationships Atlantic cod Gadus morhua, flounders) and TPNNV infects a single species (Tiger puffer Tak- Among the different species of betanodaviruses, ifugu rubripes) at an intermediate temperature amino acid sequences of RdRp protein and capsid (20 °C). The SJNNV type was initially known to protein share 87–99% and 77–100% of identity, affect a few species cultured in Japan at 20–25 °C respectively [82–98% for the complete RNA1 (Nishizawa et al. 1995; Iwamoto et al. 2000; nucleic sequence and 76–99% for the RNA2 seg- Munday et al. 2002; Toffan et al. 2016). How- ment (Okinaka & Nakai 2008)].
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