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Cloning strategy and putative genome organization of SnAdV-1 isolated from a corn snake compared to that of DAdV-1. A fragment missed by the random cloning between two Pst I clones was amplified by PCR and cloned. 

Cloning strategy and putative genome organization of SnAdV-1 isolated from a corn snake compared to that of DAdV-1. A fragment missed by the random cloning between two Pst I clones was amplified by PCR and cloned. 

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Approximately 60% of the genome of an adenovirus isolated from a corn snake (Elaphe guttata) was cloned and sequenced. The results of homology searches showed that the genes of the corn snake adenovirus (SnAdV-1) were closest to their counterparts in members of the recently proposed new genus ATADENOVIRUS: In phylogenetic analyses of the complete h...

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... was assembled ; however, the two ends and a fragment (probably larger) from the right-hand side are missing. Since the  homology search results most frequently showed the duck adenovirus 1 (DAdV-1) [syn. egg drop syndrome (EDS) virus] genes as closest relatives of the SnAdV-1 counterparts, the putative physical and genetic map presented in Fig. 1 was compared to the DAdV-1 genome. A fragment of approximately 1700 bp, located close to the left- hand end between the IVa2 protein and DNA polymerase genes and not obtained in the random cloning, was successfully amplified by PCR. The genes of the following known adenoviral proteins were identified by using the  homology search ...

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... The adenovirus of this frog (FrAdV-1) was found to be related to an unusual avian adenovirus, turkey adenovirus type 3 (TAdV-3), which is also referred to as hemorrhagic enteritis virus [15]. In recent studies, the AdV (SnAdV-1) genome isolated from the corn snake Elaphe guttata has been studied in sufficient detail [16]. The presence of AdV-like particles has been observed in several fish species; however, currently, only the gene isolate of the white sturgeon Acipenser transmontanus is available. ...
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To date, data on the presence of adenoviral receptors in fish are very limited. In the present work, we used mouse recombinant adeno-associated viral vectors (rAAV) with a calcium indicator of the latest generation GCaMP6m that are usually applied for the dorsal hippocampus of mice but were not previously used for gene delivery into fish brain. The aim of our work was to study the feasibility of transduction of rAAV in the mouse hippocampus into brain cells of juvenile chum salmon and subsequent determination of the phenotype of rAAV-labeled cells by confocal laser scanning microscopy (CLSM). Delivery of the gene in vivo was carried out by intracranial injection of a GCaMP6m-GFP-containing vector directly into the mesencephalic tegmentum region of juvenile (one-year-old) chum salmon, Oncorhynchus keta. AAV incorporation into brain cells of the juvenile chum salmon was assessed at 1 week after a single injection of the vector. AAV expression in various areas of the thalamus, pretectum, posterior-tuberal region, postcommissural region, medial and lateral regions of the tegmentum, and mesencephalic reticular formation of juvenile O. keta was evaluated using CLSM followed by immunohistochemical analysis of the localization of the neuron-specific calcium binding protein HuCD in combination with nuclear staining with DAPI. The results of the analysis showed partial colocalization of cells expressing GCaMP6m-GFP with red fluorescent HuCD protein. Thus, cells of the thalamus, posterior tuberal region, mesencephalic tegmentum, cells of the accessory visual system, mesencephalic reticular formation, hypothalamus, and postcommissural region of the mesencephalon of juvenile chum salmon expressing GCaMP6m-GFP were attributed to the neuron-specific line of chum salmon brain cells, which indicates the ability of hippocampal mammal rAAV to integrate into neurons of the central nervous system of fish with subsequent expression of viral proteins, which obviously indicates the neuronal expression of a mammalian adenoviral receptor homolog by juvenile chum salmon neurons.
... One of these genera, the Atadenovirus, was added in 2002, to include adenoviruses that were previously assigned to the genus Mastadenovirus, but varied significantly based on genomic size, structure, genes, and gene arrangement. Originally all the viruses in the genus were thought to have an A+T content bias [2,3], but with the discovery of new species in this genus, it has been shown that the A+T bias is not a consistent feature of this genus [4][5][6]. The size of sequenced atadenovirus genomes ranges between 27 and 34 kb, and all have the characteristic inverted terminal repeat (ITR) found in all adenoviruses [7]. ...
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Wild birds harbour a large number of adenoviruses that remain uncharacterised with respect to their genomic organisation, diversity, and evolution within complex ecosystems. Here, we present the first complete genome sequence of an atadenovirus from a passerine bird that is tentatively named Passerine adenovirus 1 (PaAdV-1). The PaAdV-1 genome is 39,664 bp in length, which was the longest atadenovirus to be sequenced, to the best of our knowledge, and contained 42 putative genes. Its genome organisation was characteristic of the members of genus Atadenovirus; however, the novel PaAdV-1 genome was highly divergent and showed the highest sequence similarity with psittacine adenovirus-3 (55.58%). Importantly, PaAdV-1 complete genome was deemed to contain 17 predicted novel genes that were not present in any other adenoviruses sequenced to date, with several of these predicted novel genes encoding proteins that harbour transmembrane helices. Subsequent analysis of the novel PaAdV-1 genome positioned phylogenetically to a distinct sub-clade with all others sequenced atadenoviruses and did not show any obvious close evolutionary relationship. This study concluded that the PaAdV-1 complete genome described here is not closely related to any other adenovirus isolated from avian or other natural host species and that it should be considered a separate species.
... Non-reptile atadenovirus genomes are characterised by 57.0-66.3% A + T content, whereas reptile atadenoviruses are not affected by such bias ( Farkas et al. 2002;Harrach 2008;Papp et al. 2009;Wellehan et al. 2004). Examples from other virus families are known: the influenzaviral nucleotide composition changes following a host jump from bird to mammal (Greenbaum et al. 2008), or similar differences were observed between flaviviruses or herpesviruses of different hosts ( Jenkins et al. 2001;McGeoch et al. 2006). ...
... Obviously, this composition has an effect on codon usage, thus the complete genome sequence might be under evolutionary pressure ( Shackelton et al. 2006). Still, the exact cause of this bias in atadenoviruses is unknown, but it is hypothesised that longer coevolutionary times-adaptation of the virus to the host-result in a balanced nucleotide composition ( Farkas et al. 2002;Papp et al. 2009;Wellehan et al. 2004). ...
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Viruses have been infecting their host cells since the dawn of life, and this extremely long-term coevolution gave rise to some surprising consequences for the entire tree of life. It is hypothesised that viruses might have contributed to the formation of the first cellular life form, or that even the eukaryotic cell nucleus originates from an infection by a coated virus. The continuous struggle between viruses and their hosts to maintain at least a constant fitness level led to the development of an unceasing arms race, where weapons are often shuttled between the participants. In this literature review we try to give a short insight into some general consequences or traits of virus–host coevolution, and after this we zoom in to the viral clades of adenoviruses, herpesviruses, nucleo-cytoplasmic large DNA viruses, polyomaviruses and, finally, circoviruses.
... Genome annotation was carried out with Artemis Genome Browser (Berriman and Rutherford, 2003) and CLC Main Workbench. The determination of the splicing donor and acceptor sites, as well as the cleavage signals, was accomplished manually by comparison of conserved patterns (Farkas et al., 2002;Ruzindana-Umunyana et al., 2002;Mangel and San Martin, 2014;Podgorski et al., 2016). Multiple alignments of the amino acid sequences were prepared with the Muscle program of the MEGA6 package. ...
... A schematic genetic map of the PBAdV-1 is presented in Fig. 1. The protease cleavage sites of the precursor proteins could be determined by the alignment of consensus sites (M/L/I)XGG'X (type I), (M/L/I)XGX'G (type II) and NTGW'G (type IIb) of selected adenoviruses (Farkas et al., 2002) ( Table 2 and Fig. 2). ...
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The presence of a novel adenovirus (AdV) was detected by PCR and sequencing, in the internal organs of a captive polar bear that had died in the Budapest zoo. The virus content of the samples proved to be high enough to allow for conventional Sanger sequencing on PCR-amplified genomic fragments. With this approach, the sequence of the entire genome of the putative polar bear adenovirus 1 (PBAdV-1) was obtained. Although the genome was found to be short, consisting of 27,952 base pairs merely, with a relatively balanced G + C content of 46.3%, its organisation corresponded largely to that of a typical mastadenovirus. Every genus-common gene could be identified except that of protein IX. The short E3 region of the PBAdV-1 consisted of two novel, supposedly type-specific ORFs only, whereas no homologue of any of the E3 genes, usually conserved in mastadenoviruses, such as for example that of the 12.5 K protein, were present. In the E4 region, only the highly conserved gene of the 34 K protein was found besides two novel ORFs showing no homology to any known E4 ORFs. In silico sequence analysis revealed putative splicing donor and acceptor sites in the genes of the E1A, IVa2, DNA-dependent DNA polymerase, pTP, 33 K proteins, and also of U exon protein, all being characteristic for mastadenoviruses. Phylogenetic calculations, based on various proteins, further supported that the newly-detected PBAdV is the representative of a new species within the genus Mastadenovirus, and may represent the evolutionary lineage of adenoviruses that coevolved with carnivorans.
... Notably, these latter viruses have extremely high A + T content, and are capable of causing severe diseases in certain birds or in ruminants (Benkő and Harrach, 1998). In contrast, atadenoviruses, found commonly in squamate reptiles, have a balanced G + C content and alone cause clinical disease seldom (Farkas et al., 2002). According to our hypothesis, this AdV lineage has co-evolved with the squamate reptiles and their presence in representatives of closely related hosts, or in animals classified into more distant vertebrate orders and classes, e.g. in non-squamate reptiles, different birds and multiple ruminant mammals, is the result of several ancient or more recent host switching events (Wellehan et al., 2004;Rivera et al., 2009). ...
Article
Adenoviruses are commonly found in members of almost every vertebrate lineage except fish and amphibians, from each of which only a single isolate is available as yet. In this work, the complete genomic sequence of a fish adenovirus, originating from the white sturgeon (Acipenser transmontanus), was determined and analyzed. Several exceptional features were observed including the longest hitherto known genome size (of 48,395 bp) and a strange location of the putative fiber genes resulting in an unconventional organization pattern. The left genome end contained four fiber-like genes, three of them in a tandem position on the r (rightward transcribed) strand, followed by a fourth one on the l strand. Rightward from these, the conserved adenoviral gene cassette, encompassing 16 family-common genes, was identified. In the right-hand part, amounting for >42% of the entire genome, the presence of 28 ORFs, with a coding capacity of larger than 50 amino acids, was revealed. Interestingly, most of these showed no similarity to any adenoviral genes except two ORFs, resembling slightly the parvoviral NS gene, homologues of which occur in certain avian adenoviruses. These specific traits, together with the results of phylogeny reconstructions, fully justified the separation of the white sturgeon adenovirus into the recently established new genus Ichtadenovirus. Targeted attempts to find additional adenoviruses in any other fish species were to no avail as yet. Thus the founding member, WSAdV-1 still remains the only representative of ichtadenoviruses.
... Viruses have long been known to cause morbidity and mortality throughout eukaryotes, and while much of the field of virology has been focused on human health, recent work has also begun to explore viral infections in other taxa (Davison, Wright & Harrach, 2000;Harrach, 2000;Benkő et al., 2002;Rivera et al., 2009;Kovács et al., 2010). In reptiles, pet trade husbandry has allowed for the study of viruses infecting controlled captive populations, providing to a robust understanding of several diseases in captivity (Raymond et al., 2003;Garner et al., 2008;Abbas et al., 2011;Hyndman & Shilton, 2011;Doneley, Buckle & Hulse, 2014;Bak, Yeonsook & Woo, 2018). ...
... However, host switching has also occurred. Phylogenetic study of AdVs present in deeply divergent tetrapod hosts suggest that AdVs in the genus Atadenovirus have been transferred between reptiles and birds, marsupial mammals, and ruminant mammals (Farkas et al., 2002;Farkas, Harrach & Benkő, 2008). In addition, within reptiles, distantly related taxa have been shown to harbor identical AdVs, suggesting more recent transfer of AdVs between divergent host lineages (Hyndman & Shilton, 2011;Marschang et al., 2003). ...
... We ran MrBayes for 300 million generations with one cold and three heated chains, sampling every 10,000 generations. White sturgeon AdV (Benkő et al., 2002) and Box Turtle AdV (Farkas & Gál, 2009) were selected as outgroups. After evaluation, the first 50% of trees were removed as burn-in. ...
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Adenoviruses (AdVs) infect a wide range of hosts, and they have undergone recent and ancient host transfers multiple times. In reptiles, AdVs have been found in many captive individuals, and have been implicated in morbidity and mortality in several species. Yet the pathogenicity, transmission, phylogenetic distribution, and source of AdVs in the environment are still unknown. We therefore chose to opportunistically sample deceased captive Anolis sagrei individuals that were collected from different populations in the Bahamas and the Cayman Islands, as well as fecal samples from one island population, to explore the disease dynamics and diversity of adenovirus infecting A. sagrei populations. We found that adenovirus infection was present in our captive colony at low prevalence (26%), and was likely not the primary cause of observed morbidity and mortality. Among the 10 individuals (out of 38 sampled) which tested positive for adenovirus, we identified four adenovirus clades, several of which are distantly related, despite the close relationships of the A. sagrei host populations. These results suggest that while adenovirus may not be highly prevalent in the wild, it is present at low levels across much of the range of A. sagrei . It may undergo frequent host switching across both deep and shallow host divergences.
... Adenovirus infection has been reported in snakes. [20][21][22][23][24] Raymond et al. 26 reported foci of gliosis within the CNS, and several glial and endothelial cells contained intranuclear adenovirus-like inclusion bodies in infected snakes. ...
... 1,10,15 Adenoviruses have been described as opportunistic pathogens in many animal species, especially when additional factors (particularly concurrent infections) adversely affect the health of the host; however, some types of adenovirus can have a primary pathogenic role. 1 The adenoviruses found in snakes and lizards and studied by molecular techniques all belonged to the genus Atadenovirus. 3,4,6,7,9 In chelonians, different types of adenoviral infections have been described. A systemic adenoviral infection that was associated with anorexia, lethargy, mucosal ulcerations, palatine erosions of the oral cavity, nasal and ocular discharge, and diarrhea was reported in Forsten's tortoises (Indotestudo forstenii), and the virus was shown to belong to genus Siadenovirus. ...
Article
A two-year-old female, spur-thighed tortoise (Testudo graeca) presented with poor body condition (1/5) and weakness. Fecal analysis revealed the presence of a large number of Oxyurid-like eggs, and radiographs were compatible with gastrointestinal obstruction. Despite supportive medical treatment, the animal died. At gross examination, an intestinal obstruction was confirmed. Histopathology revealed severe hyperplastic esophagitis and stomatitis with marked epithelial cytomegaly and enormous basophilic intranuclear inclusion bodies. Electron microscopy examination revealed a large number of 60-80 nm, non-enveloped, icosahedral virions arranged in crystalline arrays within nuclear inclusions of esophageal epithelial cells, morphologically compatible with adenovirus-like particles. PCR for virus identification was performed with DNA extracted from formalin-fixed paraffin-embedded tissues. A nested, consensus pan-adenovirus PCR and sequencing analysis showed a previously unidentified, novel adenovirus. According to phylogenetic calculations, it clustered to genus Atadenovirus in contrast with all other chelonian adenoviruses described to date. The present report details the pathological findings associated with an adenovirus infection restricted to the upper digestive tract.
... Infections with adenoviruses are well documented, and have been reported in all of the bony vertebrates, including squamates, from which the genus Atadenovirus likely originated. 3,5,19 Currently, there are 5 accepted genera of adenoviruses within the family Adenoviridae: Atadenovirus (predominantly in squamate hosts), Aviadenovirus (in avian hosts), Ichtadenovirus (isolated from a white sturgeon), Mastadenovirus (in mammalian hosts), and Siadenovirus (found in chelonian, amphibian, and avian species). Additionally, a sixth genus has been proposed (Testadenovirus), whose members were isolated from hosts in the order Testudines. 2 Many reptilian adenoviruses appear to have coevolved with the species they infect, leading to a fair amount of epidemiological species specificity. ...
Article
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Agamid adenovirus 1 (AgAdv-1) is a significant cause of disease in bearded dragons (Pogona sp.). Clinical manifestations of AgAdv-1 infection are variable and often nonspecific; the manifestations range from lethargy, weight loss, and inappetence, to severe enteritis, hepatitis, and sudden death. Currently, diagnosis of AgAdv-1 infection is achieved through a single published method: standard nested polymerase chain reaction (nPCR) and sequencing. Standard nPCR with sequencing provides reliable sensitivity, specificity, and validation of PCR products. However, this process is comparatively expensive, laborious, and slow. Probe hybridization, as used in a TaqMan assay, represents the best option for validating PCR products aside from the time-consuming process of sequencing. This study developed a real-time PCR (qPCR) assay using a TaqMan probe-based assay, targeting a highly conserved region of the AgAdv-1 genome. Standard curves were generated, detection results were compared with the gold standard conventional PCR and sequencing assay, and limits of detection were determined. Additionally, the qPCR assay was run on samples known to be positive for AgAdv-1 and samples known to be positive for other adenoviruses. Based on the results of these evaluations, this assay allows for a less expensive, rapid, quantitative detection of AgAdv-1 in bearded dragons. © 2015 The Author(s).
... AdVs are regularly detected in various species of lizards (Frye et al., 1994;Wellehan et al., 2004;Papp et al., 2009;Hyndman and Shilton, 2011;Ball et al., 2012Ball et al., , 2014a; in particular, central bearded dragons (Pogona vitticeps) (Kim et al., 2002;Wellehan et al., 2004;Moormann et al., 2009). They have also been described in a range of different species of snakes (Heldstab and Bestetti, 1984;Farkas et al., 2002;Marschang et al., 2003;Garner et al., 2008;Papp et al., 2009;Abbas et al., 2011). ...
... There appears to be a relative lack of species specificity among snake AdVs. Snake AdV-1 has been found in both colubrid and boid snakes (Farkas et al., 2002;Marschang et al., 2003), while snake AdV-2 has been described in viperid and colubrid snakes (Garner et al., 2008;Papp et al., 2009) and snake AdV-3 has been detected in different species of colubrid snakes (Garner et al., 2008). Thus, AdVs are not necessarily host-specific and do not always follow a co-speciation model under which host and virus phylogenies are concordant (Ascher et al., 2013). ...
... Until 2009, AdVs had been isolated only from snakes and these viruses were closely related (Jacobson et al., 1985;Juhász and Ahne, 1992;Farkas et al., 2002Farkas et al., , 2008Marschang et al., 2003). In 2009, helodermatid AdVs-1 and -2 were isolated in cell culture from different species of helodermatid lizards (H. ...