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Mccoll Khvlitreview Review of the literature on cyprinid herpesvirus 3 (CyHV-3) and its disease Kenneth A McColl Review of the literature on cyprinid herpesvirus 3 (CyHV-3) and its disease Kenneth A McColl Commonwealth Scientific and Industrial Research Organisation (CSIRO) Australian Animal Health Laboratory (AAHL) Geelong, Victoria 2013 An IA CRC Project Disclaimer: The views and opinions expressed in this report reflect those of the authors and do not necessarily reflect those of the Australian Government, Invasive Animals Ltd, or Invasive Animals Cooperative Research Centre. The material presented in this report is based on sources that are believed to be reliable. Whilst every care has been taken in the preparation of the report, it is “as is”, without warranty of any kind, to the extent permitted by law. Published by: Invasive Animals Cooperative Research Centre. Telephone: (02) 6201 2887 Facsimile: (02) 6201 2532 Email: [email protected] Internet: http://www.invasiveanimals.com Web ISBN: 978-1-921777-77-6 © Invasive Animals Ltd 2013 This work is copyright. The Copyright Act 1968 permits fair dealing for study, research, information or educational purposes. Selected passages, tables or diagrams may be reproduced for such purposes provided acknowledgement of the source is included. Major extracts of the entire document may not be reproduced by any process. The IA CRC gratefully acknowledges funding support from the Australian Government through its Cooperative Research Centres Program. This document should be cited as: McColl KA (2013). Review of the literature on cyprinid herpesvirus 3 (CyHV-3) and its disease. PestSmart Toolkit publication, Invasive Animals Cooperative Research Centre, Canberra, Australia. ii Invasive Animals CRC Contents 1. Introduction ............................................................................. 1 2. Aetiology ................................................................................. 1 3. Molecular virology ...................................................................... 2 4. Epidemiology ............................................................................ 3 4.1 Molecular epidemiology .................................................................. 3 4.2 Distribution ................................................................................. 4 4.3 Temperature susceptibility .............................................................. 5 4.4 Survival of CyHV-3 in water and sediments.......................................... 6 4.5 Transmission ................................................................................ 7 4.6 Affected species ........................................................................... 8 4.7 Susceptibility of hybrids ................................................................. 9 4.8 Age susceptibility ......................................................................... 10 4.9 Carrier Fish ................................................................................ 10 4.10 Virus activity in natural populations of carp ....................................... 11 5. Clinical signs ........................................................................... 12 6. Pathogenesis .......................................................................... 12 7. Persistence/Latency ................................................................. 13 8. Immune Response .................................................................... 15 9. Pathology............................................................................... 16 9.1 Gross Pathology ........................................................................... 16 9.2 Histopathology ............................................................................ 17 10. Differential diagnosis ................................................................ 18 11. Diagnosis ............................................................................... 19 11.1 Virus isolation ............................................................................. 19 11.2 Polymerase chain reaction / Loop mediated isothermal amplification (PCR/LAMP) ............................................................................. 20 11.3 Immunochemistry/Immunohistochemistry .......................................... 22 11.4 Electron microscopy ..................................................................... 22 11.5 Serology ..................................................................................... 22 11.6 In situ hybridisation ...................................................................... 23 11.7 Techniques for diagnosis ................................................................ 23 12. Control .................................................................................. 23 12.1 Vaccines .................................................................................... 23 12.2 Anti-viral treatments .................................................................... 24 12.3 ‘Natural’ vaccination .................................................................... 24 12.4 Crossbreeding for resistance ........................................................... 25 12.5 RNA interference (RNAI) ................................................................ 25 12.6 Control practices.......................................................................... 25 References ................................................................................. 26 Literature review: CyHV-3 iii iv Invasive Animals CRC 1. Introduction A disease in carp that was associated with high mortality and that appeared to be of viral aetiology was first described in 1997 in Germany (Bretzinger et al, 1999), and from Israel and the USA in 1998 (Hedrick et al, 2000). There is evidence that CyHV-3 was actually present in carp in the UK in 1996 (Aoki et al, 2007). It has since spread to many other countries around the world. The aetiological agent was considered to be a herpesvirus (Koi herpesvirus, KHV, now known as CyHV-3), and, in experimental infections, was associated with up to 100% mortality in both common carp (Cyprinus carpio carpio) and koi carp (C carpio koi). Furthermore, CyHV-3 affected fish of any age in either species (Hedrick et al, 2000). The disease is now considered to represent a significant threat to the important carp industry, not only in Eastern Europe (Haenen et al, 2004), but worldwide where the common carp is the world’s fourth most-farmed fish (Ronen et al, 2003). 2. Aetiology It was recommended that KHV be classified as CyHV-3 (Waltzek et al, 2005). Waltzek et al (2005) noted that herpesviruses have a double-stranded (ds) DNA genome located within an icosahedral capsid that is surrounded by a proteinaceous tegument and an outer envelope. The capsid is composed of 162 hexameric and pentameric capsomeres. Despite the recommendation for the virus to be called CyHV-3, Ilouze et al (2006) noted that the International Committee on Taxonomy of Viruses had not yet officially named the virus. Unofficially, it has been called KHV, Carp interstitial Nephritis and Gill Necrosis Virus (CNGV), and koi herpes-like virus. However, they claimed that, while the virus looked like a herpesvirus, the large genome (larger than any known herpesvirus apart from the other cyprinid herpesviruses), and the presence of genes that encode peptides that are found in other large dsDNA viruses (but not herpesviruses), suggested that CyHV-3 may belong to a novel virus group. There was a suggestion that CyHV-3 can be differentiated from another herpesvirus of cyprinids, Cyprinid herpesvirus (CHV, also known as CyHV-1), based on the age of affected fish (Gilad et al, 2003; 2004), ie, CyHV-3 affects fish of any age, while CyHV-1, that can also affect both subspecies of carp, generally only kills young fish upto 2 months old (Sano et al, 1985a,b; 1991). It should be noted that CyHV-1 is also the cause of “carp pox” (Sano et al, 1985a,b), and upto 60% of fry surviving the acute form of the disease may develop epidermal tumours/hyperplastic papillomas (Sano et al, 1991; Hedrick et al, 1990). CyHV-1-like lesions are seen regularly in Australia, especially in koi carp (Fran Stephens, pers comm). This disease has been present for many years, although there appear to be no formal reports of its presence. In general, CyHV-3 and CyHV-1 can be differentiated by (1) differences in cell-line specificity (CyHV-1 grows on EPC and FHM cyprinid cell lines); (2) type of cytopathic effect (CPE) in cell culture; (3) antigenic properties. Hedrick et al (2000) demonstrated that anti-CyHV-1 Literature review: CyHV-3 1 antiserum recognized CyHV-1-infected cells, but not CyHV-3-infected cells; and, (4) timing of lesions (CyHV-1 lesions occur when water temperatures are declining [Hedrick et al, 1990]). Note that goldfish (Carassius [Cyprinus] auratus), which are also cyprinids, are not affected by either of these viruses. Hedrick et al (2000) and Gilad et al (2004) also mentioned another herpesvirus of cyprinid fish, goldfish haematopoietic necrosis virus (GHNV also known as CyHV-2). CyHV-2-lesions have been detected on a number of occasions, since about 2001, in goldfish in Australia (Stephens et al, 2004). Neither common carp nor koi carp appear to be susceptible to CyHV-2 (Hedrick et al, 2006). 3. Molecular virology Waltzek and Hedrick (2004) noted that the complete genome sequence was available for CyHV-3. Based on the sequences of four complete genes (two encoding proteins involved in DNA replication,
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