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Avian Pox Virus |   373

          fowlpox and candidiasis in backyard chickens with unusual pox   highly pathogenic fowlpox virus with high mortality. The authors
          lesions in the bursa of Fabricius (BF) confirmed by PCR amplifi-  believed that the virulence was associated with the integration of
          cation of 4b and TK genes. Pox lesions were not observed in the   REV sequences in the FWPV genome. The flock had not received
          internal organs except for the BF.                    any vaccination. When  immune responses of  fowlpox  vaccine
                                                                strains and field isolates were compared (Wang et al., 2006),
                                                                CMI responses were significantly depressed in chickens inocu-
          Role of REV in the fowlpox virus genome               lated with field isolates which appeared in the first 3 weeks post
          The FWPV genome contains an ORF that encodes for a putative   infection. The field strains of FWPV contained integrated REV
          protein with similarity to the protein encoded by Marek’s disease   provirus while vaccine strains carry only REV LTR sequences.
          virus  and  fowl  adenovirus,  the  presence  of  this  homologue  in   The results of this study further suggest that virulent field isolates
          three different DNA viruses of birds suggests its important role   FWPV with full-length REV have immunosuppressive effects
          in avian host range function. Interestingly, a natural dual viral   when inoculated into young chickens.
          infection of trachea by FWPV and herpesvirus has been reported
          previously (Tripathy et al., 1975). In this case presence of herpes
          virus in the nucleus and poxvirus in cytoplasm of the same cell   Prevention
          was demonstrated by histopathology and electron microscopy. As   Live virus vaccines of FWPV and pigeonpox virus origin are
          homologues of FWPV open reading frames were detected in the   used in domestic poultry in areas where the disease is endemic.
          genome of Marek’s disease virus (Brunovskis and Velicer, 1995),   Chicken embryos or avian cell cultures are used for propagation
          the likelihood of exchange of genetic material from one virus to   of vaccine viruses.
          another and emergence of a genetically and antigenically differ-  The vaccines are administered by wing-web stab method. Vac-
          ent virus is possible. In this regard, integration of full-length REV   cinated birds are examined after 7–10 days, for swelling of the skin
          in majority of the field isolates of the FWPV genome indicates   ‘takes’. Attenuated cell-culture vaccine can be used on chicks as
          an event of ‘natural genetic engineering’ in viruses. Both experi-  early as day old. Although the vaccine is administered to 4-week-
          mental and field observation (Tripathy et al., 1998; Singh et al.,   old chickens, and again at 1–2 months prior to egg production,
          2000) indicate fowlpox as an emerging/re-emerging disease. Kim   the practice may vary depending upon the epizootiology of the
          and Tripathy (2001) observed that an isolate of fowlpox virus   disease in a particular operation and area. The chickens may be
          lyophilized in 1949 had full-length REV insertion in its genome   revaccinated if held for more than one laying cycle. Experimental
          indicating that this REV insertion happened long ago. While the   studies show that FWPV vaccines given in ovo provide protection
          information on the origin and date of isolation of fowlpox virus   against the disease. A cell-culture FWPV vaccine provides protec-
          strain were not available, it is assumed that the event occurred   tion against FWPV in day-old turkeys (Sarma et al., 2015).
          sometime in 1930–1940 or earlier. Phylogenetic and historical
          evidences (Niewiadomska and Gifford, 2013) suggest that REVs   Recombinant FWPV and CNPV vaccines
          originated as mammalian retroviruses which were accidentally   A large variety of genes encoding antigenic proteins of specific
          introduced into avian hosts in the late 1930s, during experimen-  pathogens have been inserted into the genome of FWPV and
          tal studies, and subsequently integrated into the FWPV genomes,   CNPV. Because of its large genome size with several non-essential
          generating recombinant DNA viruses that now circulate in wild   loci, genes from more than one pathogen can be inserted into its
          birds and poultry. REV has been associated with immunosup-  genome to create a polyvalent vaccine (Boyle and Heine, 1993;
          pression and tumour formation.                        Boyle et al., 2004).
            Only variable-length, REV LTR remnants are present in the   Several non-essential regions, including some in the terminal
          genome of all FWPV vaccine strains (Moore et al., 2000). These   inverted repeats, have been identified in the FWPV genome and
          remnants are also retained, presumably after the loss of the REV   have been used in the generation of recombinant viruses. Thymi-
          provirus, by a minor proportion of each field strain population.  dine Kinase (TK) gene has been one of the common loci where
            Isolation of majority of FWPV strains with intact provirus   foreign genes have been inserted. Since TK is associated with
          copy of REV genome from chicken flocks which had been vac-  virulence and is non-essential for virus replication, the recom-
          cinated with fowlpox and pigeonpox virus vaccines is indicative   binant FWPV are less virulent than parent FWPV (Beard et al.,
          of emergence of strains which are antigenically, and biologically   1991). Pox virus promoters are relatively conserved and, thus, are
          different and current vaccines do not provide adequate protec-  recognized by heterologous pox viruses. Initially, therefore, vac-
          tion against them.                                    cinia virus promoters were used in lieu of FWPV transcription
            Both immunosuppression and tumour formation by REV   regulatory elements in creating recombinant FWPV. Although
          containing FWPV are supported by field observations. For exam-  homologous FWPV promoters have since been identified
          ple, visceral lymphomas were observed in chickens infected with   (Srinivasan et al., 2006) and a synthetic, early late transcriptional
          fowlpox virus containing REV by Koo et al. (2013). Since sera of   regulatory element has been used, 2 vaccinia virus promoters, the
          affected birds were positive for REV specific antibodies but not   early-late P7.5 and the late P11, have been used for the construc-
          for avian leukosis virus, the authors concluded that the lympho-  tion of recombinant avian pox viruses (Beard et al., 1991). Several
          mas were caused by the REV containing FWPV field strain. In   homologous FWPV promoters, including a bidirectional one,
          another observation, Zhao et al. (2014) reported an outbreak of   have been evaluated. Because some of these promoters appear
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