Family Alloherpesviridae Research Articles

ТАКСОНОМІЧНА ХАРАКТЕРИСТИКА ДНК-ГЕНОМНИХ ВІРУСІВ ХРЕБЕТНИХ ТВАРИН І ЛЮДИНИ

Presented modern taxonomy and nomenclature of the DNA-genomic of viruses of vertebrates animals and humans in accordance with the information ICTV release 2015 (ratification 2016). Described the basic criteria for the classification of viruses: characteristics of the viral genome, the mechanism of replication and virions morphology. Viruses of vertebrates (1120 species) consist of 4 orders, 34 families (12 – DNA-genomic, 22 – RNA-genomic), 11 subfamilies and 219 genera. DNA-genomic viruses of vertebrates (546 species) classified in 1 orders, 12 families, 5 subfamilies and 113 genera. The order Herpesvirales has united family Herpesviridae and Alloherpesviridae. Family Poxviridae, Iridoviridae and Parvoviridae, except of viruses of vertebrates, contain viruses of insects. Described the taxa of viruses:family, subfamily, genera, species. Characterized the basic taxonomic features of DNA-genomic viruses of vertebrates: the shape, size and structure of virions – the presence of outer membrane lipoprotein, capsid symmetry type, the structure of the viral DN and the number of proteins. The attention is focused on the features of reproduction of viruses. The replication of majority DNA-genomic viruses of vertebrates occur in the nucleus of cells, except for members of families Poxviridae and Asfarviridae, which are replicate in the cytoplasm. Yield virions is done due to destruction of cells or budding through the cell membrane (sometimes in combination with exocytosis) depending on the structural organization of the virus.

Complete genome sequence and architecture of crucian carp Carassius auratus herpesvirus (CaHV).

Crucian carp Carassius auratus herpesvirus (CaHV) was isolated from diseased crucian carp with acute gill hemorrhages and high mortality. The CaHV genome was sequenced and analyzed. The data showed that it consists of 275,348 bp and contains 150 predicted ORFs. The architecture of the CaHV genome differs from those of four cyprinid herpesviruses (CyHV1, CyHV2, SY-C1, CyHV3), with insertions, deletions and the absence of a terminal direct repeat. Phylogenetic analysis of the DNA polymerase sequences of 17 strains of Herpesvirales members, and the concatenated 12 core ORFs from 10 strains of alloherpesviruses showed that CaHV clustered together with members of the genus Cyprinivirus, family Alloherpesviridae.

A Quantitative Polymerase Chain Reaction Assay for the Detection and Quantification of Epizootic Epitheliotropic Disease Virus (EEDV; Salmonid Herpesvirus 3).

Epizootic epitheliotropic disease virus (EEDV; salmonid herpesvirus [SalHV3]; family Alloherpesviridae) causes a systemic disease of juvenile and yearling Lake Trout Salvelinus namaycush. No cell lines are currently available for the culture and propagation of EEDV, so primary diagnosis is limited to PCR and electron microscopy. To better understand the pervasiveness of EEDV (carrier or latent state of infection) in domesticated and wild Lake Trout populations, we developed a sensitive TaqMan quantitative PCR (qPCR) assay to detect the presence of the EEDV terminase gene in Lake Trout tissues. This assay was able to detect a linear standard curve over nine logs of plasmid dilution and was sensitive enough to detect single-digit copies of EEDV. The efficiency of the PCR assay was 99.4 ± 0.06% (mean ± SD), with a 95% confidence limit of 0.0296 (R(2) = 0.994). Methods were successfully applied to collect preliminary data from a number of species and water bodies in the states of Pennsylvania, New York, and Vermont, indicating that EEDV is more common in wild fish than previously known. In addition, through the development of this qPCR assay, we detected EEDV in a new salmonid species, the Cisco Coregonus artedi. The qPCR assay was unexpectedly able to detect two additional herpesviruses, the Atlantic Salmon papillomatosis virus (ASPV; SalHV4) and the Namaycush herpesvirus (NamHV; SalHV5), which both share high sequence identity with the EEDV terminase gene. With these unexpected findings, we subsequently designed three primer sets to confirm initial TaqMan qPCR assay positives and to differentiate among EEDV, ASPV, and NamHV by detecting the glycoprotein genes via SYBR Green qPCR. Received April 20, 2015; accepted November 10, 2015.

Initial Detection and Molecular Characterization of Namaycush Herpesvirus (Salmonid Herpesvirus 5) in Lake Trout.

A novel herpesvirus was found by molecular methods in samples of Lake Trout Salvelinus namaycush from Lake Erie, Pennsylvania, and Lake Ontario, Keuka Lake, and Lake Otsego, New York. Based on PCR amplification and partial sequencing of polymerase, terminase, and glycoprotein genes, a number of isolates were identified as a novel virus, which we have named Namaycush herpesvirus (NamHV) salmonid herpesvirus 5 (SalHV5). Phylogenetic analyses of three NamHV genes indicated strong clustering with other members of the genus Salmonivirus, placing these isolates into family Alloherpesviridae. The NamHV isolates were identical in the three partially sequenced genes; however, they varied from other salmonid herpesviruses in nucleotide sequence identity. In all three of the genes sequenced, NamHV shared the highest sequence identity with Atlantic Salmon papillomatosis virus (ASPV; SalHV4) isolated from Atlantic Salmon Salmo salar in northern Europe, including northwestern Russia. These results lead one to believe that NamHV and ASPV have a common ancestor that may have made a relatively recent host jump from Atlantic Salmon to Lake Trout or vice versa. Partial nucleotide sequence comparisons between NamHV and ASPV for the polymerase and glycoprotein genes differ by >5% and >10%, respectively. Additional nucleotide sequence comparisons between NamHV and epizootic epitheliotropic disease virus (EEDV/SalHV3) in the terminase, glycoprotein, and polymerase genes differ by >5%, >20%, and >10%, respectively. Thus, NamHV and EEDV may be occupying discrete ecological niches in Lake Trout. Even though NamHV shared the highest genetic identity with ASPV, each of these viruses has a separate host species, which also implies speciation. Additionally, NamHV has been detected over the last 4 years in four separate water bodies across two states, which suggests that NamHV is a distinct, naturally replicating lineage. This, in combination with a divergence in nucleotide sequence from EEDV, indicates that NamHV is a new species in the genus Salmonivirus. Received April 20, 2015; accepted October 11, 2015.

Is There Any Species Specificity in Infections with Aquatic Animal Herpesviruses?–The Koi Herpesvirus (KHV): An Alloherpesvirus Model

Most diseases induced by herpesviruses are host-specific; however, exceptions exist within the family Alloherpesviridae. Most members of the Alloherpesviridae are detected in at least two different species, with and without clinical signs of a disease. In the current study the Koi herpesvirus (KHV) was used as a model member of the Alloherpesviridae and rainbow trout as a model salmonid host, which were infected with KHV by immersion. KHV was detected using direct methods (qPCR and semi-nested PCR) and indirect (enzyme-linked immunosorbant assay; ELISA, serum neutralization test; SNT). The non-koi herpesvirus disease (KHVD)-susceptible salmonid fish were demonstrated to transfer KHV to naive carp at two different temperatures including a temperature most suitable for the salmonid (15°C) and cyprinid (20°C). At 20°C KHVD was induced in carp cohabitated with infected trout. KHV was also detected virologically and serologically at the end of the experiment in both rainbow trout and carp.

Molecular detection of a putatively novel cyprinid herpesvirus in sichel (Pelecus cultratus) during a mass mortality event in Hungary.

In the early summer of 2014, mass mortality of sichel (Pelecus cultratus) was observed in Lake Balaton, Hungary. Histological examination revealed degenerative changes within the tubular epithelium, mainly in the distal tubules and collecting ducts in the kidneys and multifocal vacuolisation in the brain stem and cerebellum. Routine molecular investigations showed the presence of the DNA of an unknown alloherpesvirus in some specimens. Subsequently, three genes of the putative herpesviral genome (DNA polymerase, terminase, and helicase) were amplified and partially sequenced. A phylogenetic tree reconstruction based on the concatenated sequence of these three conserved genes implied that the virus belongs to the genus Cyprinivirus within the family Alloherpesviridae. The sequences of the sichel herpesvirus differ markedly from those of the cypriniviruses CyHV-1, CyHV-2 and CyHV-3, putatively representing a fifth species in the genus.

Atlantic salmon papillomatosis in Russia and molecular characterization of the associated herpesvirus

Papillomatosis of Atlantic salmon Salmo salar has been reported for decades in Russia, Scandinavia and Scotland. The disease is typically benign although heavy losses have occasionally been reported. A herpesviral etiology has been suggested based on ultrastructural evidence; however, the virus has not been isolated or genetically characterized. In this study, we provide the first viral sequences detected in the papillomas from diseased Russian Atlantic salmon. Phylogenetic analyses, based on the partial sequences of the herpesviral polymerase and terminase genes, supported the virus as a novel member of the genus Salmonivirus within the family Alloherpesviridae. The sequences of the Atlantic salmon papillomatosis virus differ markedly from those of the 3 known salmoniviruses; therefore, the authors propose the species designation Salmonid herpesvirus 4 to be considered for approval by the International Committee on Taxonomy of Viruses.

IL-10 encoded by viruses: a remarkable example of independent acquisition of a cellular gene by viruses and its subsequent evolution in the viral genome

Many viruses have evolved strategies to deregulate the host immune system. These strategies include mechanisms to subvert or recruit the host cytokine network. IL-10 is a pleiotropic cytokine that has both immunostimulatory and immunosuppressive properties. However, its key features relate mainly to its capacity to exert potent immunosuppressive effects. Several viruses have been shown to upregulate the expression of cellular IL-10 (cIL-10) with, in some cases, enhancement of infection by suppression of immune functions. Other viruses encode functional orthologues of cIL-10, called viral IL-10s (vIL-10s). The present review is devoted to these virokines. To date, vIL-10 orthologues have been reported for 12 members of the family Herpesviridae, two members of the family Alloherpesviridae and seven members of the family Poxviridae. Study of vIL-10s demonstrated several interesting aspects on the origin and the evolution of these viral genes, e.g. the existence of multiple (potentially up to nine) independent gene acquisition events at different times during evolution, viral gene acquisition resulting from recombination with cellular genomic DNA or cDNA derived from cellular mRNA and the evolution of cellular sequence in the viral genome to restrict the biological activities of the viral orthologues to those beneficial for the virus life cycle. Here, various aspects of the vIL-10s described to date are reviewed, including their genetic organization, protein structure, origin, evolution, biological properties and potential in applied research.

Cyprinid herpesvirus 3: an interesting virus for applied and fundamental research

Cyprinid herpesvirus 3 (CyHV-3), a member of the family Alloherpesviridae is the causative agent of a lethal, highly contagious and notifiable disease in common and koi carp. The economic importance of common and koi carp industries together with the rapid spread of CyHV-3 worldwide, explain why this virus became soon after its isolation in the 1990s a subject of applied research. In addition to its economic importance, an increasing number of fundamental studies demonstrated that CyHV-3 is an original and interesting subject for fundamental research. In this review, we summarized recent advances in CyHV-3 research with a special interest for studies related to host-virus interactions.

CyHV-3: the third cyprinid herpesvirus

Common carp (including ornamental koi carp) Cyprinus carpio L. are ecologically and economically important freshwater fish in Europe and Asia. C. carpio have recently been endangered by a third cyprinid herpesvirus, known as cyprinid herpesvirus-3 (CyHV-3), the etiological agent of koi herpesvirus disease (KHVD), which causes significant morbidity and mortality in koi and common carp. Clinical and pathological signs include epidermal abrasions, excess mucus production, necrosis of gill and internal organs, and lethargy. KHVD has decimated major carp populations in Israel, Indonesia, Taiwan, Japan, Germany, Canada, and the USA, and has been listed as a notifiable disease in Germany since 2005, and by the World Organisation for Animal Health since 2007. KHVD is exacerbated in aquaculture because of the relatively high host stocking density, and CyHV-3 may be concentrated by filter-feeding aquatic organisms. CyHV-3 is taxonomically grouped within the family Alloherpesviridae, can be propagated in a number of cell lines, and is active at a temperature range of 15 to 28°C. Three isolates originating from Japan (KHV-J), USA (KHV-U), and Israel (KHV-I) have been sequenced. CyHV-3 has a 295 kb genome with 156 unique open reading frames and replicates in the cell nucleus, and mature viral particles are 170 to 200 nm in diameter. CyHV-3 can be detected by multiple PCR-based methods and by enzyme-linked immunosorbent assay. Several modes of immunization have been developed for KHVD; however, fish immunized with either vaccine or wild-type virus may become carriers for CyHV-3. There is no current treatment for KHVD.

The IL-10 homologue encoded by cyprinid herpesvirus 3 is essential neither for viral replication in vitro nor for virulence in vivo

Cyprinid herpesvirus 3 (CyHV-3), a member of the family Alloherpesviridae, is the causative agent of a lethal disease in common and koi carp. CyHV-3 ORF134 encodes an interleukin-10 (IL-10) homologue. The present study was devoted to this ORF. Transcriptomic analyses revealed that ORF134 is expressed as a spliced gene belonging to the early-late class. Proteomic analyses of CyHV-3 infected cell supernatant demonstrated that the ORF134 expression product is one of the most abundant proteins of the CyHV-3 secretome. To investigate the role of ORF134 in viral replication in vitro and in virulence in vivo, a deleted strain and a derived revertant strain were produced using BAC cloning technologies. The recombinant ORF134 deleted strain replicated in vitro comparably to the parental and the revertant strains. Infection of fish by immersion in water containing the virus induced comparable CyHV-3 disease for the three virus genotypes tested (wild type, deleted and revertant). Quantification of viral DNA by real time TaqMan PCR (in the gills and the kidney) and analysis of carp cytokine expression (in the spleen) by RT-qPCR at different times post-infection did not revealed any significant difference between the groups of fish infected with the three virus genotypes. Similarly, histological examination of the gills and the kidney of infected fish revealed no significant differences between fish infected with ORF134 deleted virus versus fish infected with the control parental or revertant strains. All together, the results of the present study demonstrate that the IL-10 homologue encoded by CyHV-3 is essential neither for viral replication in vitro nor for virulence in common carp.

Koi Herpes Virus: A Review and Risk Assessment of Indian Aquaculture

Common carp (Cyprinus carpio) is a widely cultivated freshwater fish for human consumption, while koi carp, is a farmed colored sub species of common carp used for ornamental purposes. Since 1998, both common carp and koi carp are severely affected by a viral disease called as Koi herpes virus disease (KHVD). This disease is caused by Koi herpes virus (KHV), also known as cyprinid herpes virus-3. The virus causes interstitial nephritis and gill necrosis in carps, so it is also termed as carp interstitial nephritis and gill necrosis virus. KHV is a double stranded icosahedral DNA virus belonging to family Alloherpesviridae, with a genome size of 295kbp, larger than any member of Herpesviridae. The viral genome encodes 156 potential protein coding open reading frames. Each virion consists of forty structural proteins, which are classified as capsid (3), envelope (13), tegument (2) and unclassified (22) structural proteins. Diagnosis of KHVD is mainly based on detection of viral DNA by polymerase chain reaction amplification using specific primers or loop mediated isothermal amplification. Temperature dependent latent infection is unique to KHV; and carrier fish are often not detected, thereby possibly resulting in spread of this pathogen to newer areas. The disease is now known to occur in, or has been recorded from at least 26different countries of the world. Fortunately, KHVD has not been reported from India or from Indian major carps. To monitor the disease status of the country, a total of 254 fish samples collected from different parts of India were screened by PCR for the presence of KHV. None of the tested samples were found to be positive for KHV. These results demonstrate that tested samples from different parts of India were apparently free from KHV. Preliminary risk assessment of KHV suggest that in the event of unrestricted importation of koi carps into our country, there is a higher probability of risk to aquaculture as compared to natural waters. So there is strong need to develop diagnostic capabilities and launch surveillance programmes for KHV in India.

Fish herpesvirus diseases: a short review of current knowledge

Fish herpesviruses can cause significant economic losses in aquaculture, and some of these viruses are oncogenic. The virion morphology and genome organization of fish herpesviruses are generally similar to those of higher vertebrates, but the phylogenetic connections between herpesvirus families are tenuous. In accordance with new taxonomy, fish herpesviruses belong to the family Alloherpesviridae in the order Herpesvirales. Fish herpesviruses can induce diseases ranging from mild, inapparent infections to serious ones that cause mass mortality. The aim of this work was to summarize the present knowledge about fish herpesvirus diseases.

Herpesviruses that infect fish.

Herpesviruses are host specific pathogens that are widespread among vertebrates. Genome sequence data demonstrate that most herpesviruses of fish and amphibians are grouped together (family Alloherpesviridae) and are distantly related to herpesviruses of reptiles, birds and mammals (family Herpesviridae). Yet, many of the biological processes of members of the order Herpesvirales are similar. Among the conserved characteristics are the virion structure, replication process, the ability to establish long term latency and the manipulation of the host immune response. Many of the similar processes may be due to convergent evolution. This overview of identified herpesviruses of fish discusses the diseases that alloherpesviruses cause, the biology of these viruses and the host-pathogen interactions. Much of our knowledge on the biology of Alloherpesvirdae is derived from research with two species: Ictalurid herpesvirus 1 (channel catfish virus) and Cyprinid herpesvirus 3 (koi herpesvirus).

Partial genome characterization of acipenserid herpesvirus 2: taxonomical proposal for the demarcation of three subfamilies in Alloherpesviridae

Sequencing of approximately one half of the genome of acipenserid herpesvirus 2 (AciHV-2), which is a member of the genus Ictalurivirus in the family Alloherpesviridae, revealed that the gene organization is very similar to that of ictalurid herpesvirus 1 (IcHV-1), the founder member of the genus. The sequenced region encodes the AciHV-2 homologues of IcHV-1 ORF24 to ORF69. It contains 46 predicted protein-coding regions, including 12 that seem to have a homologue in every alloherpesvirus genome sequenced to date. Phylogenetic tree reconstruction, based on the concatenated sequence of these conserved genes, implied that the family Alloherpesviridae is composed of three major clades and could be subdivided into three subfamilies.

Comparative Analysis of a Conserved Gene Block from the Genome of the Members of the Genus Ictalurivirus

Objective: Partial genome sequences were determined and subjected to comparative analyses from two fish herpesviruses (HVs). Acipenserid (Aci) HV-2, originating from the white sturgeon (Acipenser transmontanus), and ictalurid (Ic) HV-2, isolated from the black bullhead (Ameiurus melas), are recently approved species of the genus Ictalurivirus of the family Alloherpesviridae. Methods: An almost 8,000-base-pair fragment, spanning between the genes of the DNA polymerase and the ATPase subunit of the terminase, was sequenced from each virus. Results: The size, position and orientation of 2 partial and 3 full open reading frames, contained in the studied genome fragment, proved to be similar to their counterparts in IcHV-1, the type species of the genus Ictalurivirus. Thus, a well-conserved genus-specific gene block was identified. In the members of two other genera (Cyprinivirus and Batrachovirus) of the family Alloherpesviridae, no such gene block could be found; the location and orientation of the homologous genes showed significant divergence. Conclusion: The results of phylogenetic calculations were in good agreement with the genome arrangements inasmuch as AciHV-2, IcHV-1 and -2 are monophyletic and separated from the lineages of the other two genera. The new sequence enabled the inclusion of a hitherto unassigned HV, that of the Australian pilchard, into a phylogenetic calculation.

The outbreak of carp disease caused by CyHV‐3 as a model for new emerging viral diseases in aquaculture: a review

AbstractAquacultured koi and common carp (Cyprinus carpio) are intensively bred for ornamental purposes and for human consumption worldwide. The carp and koi farming industries have suffered enormous economic losses over the past decade due to an epizootic disease caused by Cyprinus herpesvirus‐3 (CyHV‐3) also known as koi herpesvirus and carp interstitial nephritis gill necrosis virus. CyHV‐3 is a large dsDNA virus, morphologically similar to herpesviruses, yet contains genetic elements similar to those of pox, irido‐ and herpesviruses. Considering the phylogenic distance between CyHV‐3 and higher vertebrate herpesviruses, CyHV‐3 represents the prototype of viruses assigned to the novel family Alloherpesviridae. Although emergence of a new virus rarely initiates a pandemic so severe that it reduces the life expectancy of a population, CyHV‐3 is exceptional because of its enormous impact on the world carp population. High population density is the major contributing factor to the epizootic disease caused by CyHV‐3.

Complete genome sequence and taxonomic position of anguillid herpesvirus 1

Eel herpesvirus or anguillid herpesvirus 1 (AngHV1) frequently causes disease in freshwater eels. The complete genome sequence of AngHV1 and its taxonomic position within the family Alloherpesviridae were determined. Shotgun sequencing revealed a 249 kbp genome including an 11 kbp terminal direct repeat that contains 7 of the 136 predicted protein-coding open reading frames. Twelve of these genes are conserved among other members of the family Alloherpesviridae and another 28 genes have clear homologues in cyprinid herpesvirus 3. Phylogenetic analyses based on amino acid sequences of five conserved genes, including the ATPase subunit of the terminase, confirm the position of AngHV1 within the family Alloherpesviridae, where it is most closely related to the cyprinid herpesviruses. Our analyses support a recent proposal to subdivide the family Alloherpesviridae into two sister clades, one containing AngHV1 and the cyprinid herpesviruses and the other containing Ictalurid herpesvirus 1 and the ranid herpesviruses.

Phylogenetic relationships in the family Alloherpesviridae

Phylogenetic relationships among herpesviruses (HVs) of mammals, birds, and reptiles have been studied extensively, whereas those among other HVs are relatively unexplored. We have reconstructed the phylogenetic relationships among 13 fish and amphibian HVs using maximum likelihood and Bayesian analyses of amino acid sequences predicted from parts of the DNA polymerase and terminase genes. The relationships among 6 of these viruses were confirmed using the partial DNA polymerase data plus the complete sequences of the terminase, helicase, and triplex protein genes; the position of these viruses among all other sequenced HVs was also investigated using the complete terminase gene. The results established the monophyly of the fish and amphibian HVs (Alloherpesviridae) separate from the HVs of mammals, birds, and reptiles (Herpesviridae) and the single recognized HV of bivalve mollusks (Malacoherpesviridae) in the order Herpesvirales. Two major clades in the family Alloherpesviridae were recognized: one consisting of viruses from cyprinid and anguillid hosts and the other of viruses from ictalurid, salmonid, acipenserid, and ranid hosts. A comparison of virus and host phylogenies suggested that closely related HVs in this family may have coevolved with their hosts, whereas significant codiversification was not apparent for the more distantly related viruses.

The order Herpesvirales

The taxonomy of herpesviruses has been updated by the International Committee on Taxonomy of Viruses (ICTV). The former family Herpesviridae has been split into three families, which have been incorporated into the new order Herpesvirales. The revised family Herpesviridae retains the mammal, bird and reptile viruses, the new family Alloherpesviridae incorporates the fish and frog viruses, and the new family Malacoherpesviridae contains a bivalve virus. Three new genera have been created in the family Herpesviridae, namely Proboscivirus in the subfamily Betaherpesvirinae and Macavirus and Percavirus in the subfamily Gammaherpesvirinae. These genera have been formed by the transfer of species from established genera and the erection of new species, and other new species have been added to some of the established genera. In addition, the names of some nonhuman primate virus species have been changed. The family Alloherpesviridae has been populated by transfer of the genus Ictalurivirus and addition of the new species Cyprinid herpesvirus 3. The family Malacoherpesviridae incorporates the new genus Ostreavirus containing the new species Ostreid herpesvirus 1.