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202  |  Kibenge et al.

          al. (2003) reported an experimental in ovo vaccination method   (Lu et al., 2011). Avian orthoreoviruses were genetically char-
          for commercial poultry.                               acterized by sequencing their σC proteins, into four genotypes
            Current  commercial  vaccine  strains  are antigenically  and   (Goldenberg et al., 2010), and the σC proteins of the currently
          serologically distinct from circulating variant field viruses associ-  used vaccine strains were found to differ from those of most field
          ated with clinical disease (Goldenberg et al., 2010; Sellers, 2017).   isolates (Vasserman et al., 2004; Goldenberg et al., 2010). Since
          Most of the orthoreovirus vaccines have been developed from   σC is the major protein against which neutralizing antibodies are
          various passage levels of avian orthoreovirus S1133 (Huang et   produced, this mismatch might lead to vaccination inefficiency
          al., 1987), 2177, 2408 or 1733 (Davis et al., 2012). Giambrone   or failure. Lublin et al. (2011) tested the efficacy of a tetra-valent
          et al. (1992) showed that S1133 was effective in the prevention   inactivated vaccine consisting of a mixture of prototypes of the
          of 2408 and CO8 enteric orthoreovirus infections when given   four defined genotypic groups of avian orthoreovirus and found
          by coarse spray to one day old SPF broiler chickens. Since 2012,   that the vaccine was able to prevent disease and confer broad
          there has been a dramatic increase in the number of tenosynovitis   protection against field isolates. Also, vaccinations with avian
          disease outbreaks in commercial poultry (Sellers, 2017). Despite   orthoreovirus prototypes from all genotypes efficiently protected
          vaccination, a high percentage of flocks have been infected with   against other members of their respective genotypes (Lublin et
          orthoreovirus, and the limited effectiveness of available vaccines   al., 2011).
          is likely due to the existence of many viral variants in the com-  Although orthoreoviruses are difficult to eliminate from the
          mercial poultry population (Lublin et al., 2011). Giambrone and   environment because of their resistance to chemical and physical
          Solano (1988) found all of the most common vaccine strains   disinfectants, decontamination of a poultry facility following total
          (S1133, 81-5, 2408, 1733 and UMI 203) belong to a single   removal of infected birds and prior to reintroduction of a new
          serotype by virus-neutralization test and ELISA, explaining their   flock prevents infection of the new population (Stott, 1999). Pro-
          inability to protect against field virus isolates belonging to other   tection of commercial poultry flocks from wild birds by providing
          serotypes. Other possible reasons for vaccination failure include   high level biosecurity, quarantine of birds to be introduced to the
          improper vaccination of pullets, decreased maternal antibody   flock, monitoring of the flock for signs of illness, serological test-
          titre in older hens, and the necessity for adequate induction of   ing and vaccination for specific diseases, and good sanitation and
          cell-mediated immunity (CMI) induced by live vaccines (Giam-  hygiene practices are all critical methods to prevent the spread
          brone et al., 1992). Subcutaneous vaccination at 1 day of age is   of avian diseases between wild birds and commercial poultry
          efficacious for induction of sufficient CMI to prevent orthoreo-  (Hollmén and Docherty, 2007). In addition, the quick removal
          virus infections (van der Heide et al., 1983). However, when   of carcasses during a wild bird die-off reduces contamination of
          orthoreovirus vaccines are combined with turkey herpesvirus   the environment and the possibility of disease transmission to
          vaccine and given subcutaneously, interference may occur, which   other avian species and/or commercial poultry (Hollmén and
          results in the reduced efficacy of both products (Giambrone and   Docherty, 2007).
          Hathcock, 1991). In areas where Marek’s disease virus exposure
          is high, increased Marek’s disease has occurred when orthoreo-
          virus and Marek’s disease vaccines were combined and given at   Perspectives
          1 day of age. Stott (1999) noted, despite in vitro cross reaction   The host range of avian reoviruses includes all domestic and
          between strains of orthoreovirus,  in  vivo  vaccination ensures   exotic or semi-domesticated poultry (chickens, turkeys, ducks,
          protection only against the homologous virus. Producers now   geese,  pigeon, quail) and  a wide range  of  free-ranging  (wild)
          use autogenous inactivated vaccines, however, identification and   birds. By their very nature, avian reoviruses have most frequently
          selection of field isolates for such use can be difficult especially   been found in avian species that were clinically normal. The
          when multiple reoviruses including new genetic variants are co-  most common and readily diagnosed disease presentation is
          circulating among flocks (Sellers, 2017).             viral arthritis/tenosynovitis in commercial broiler chickens and
            As conventional attenuated and inactivated orthoreovirus   turkeys, and enteric disease and neurological disease in wild
          vaccines are not totally efficacious (van Loon et al., 2002), new   birds. Avian reoviruses belong to the genus Orthoreovirus, one of
          approaches for vaccine production have been attempted. Vasser-  15 recognized genera in the family Reoviridae, but distinct from
          man et al. (2004) demonstrated subcutaneous injection of σC   mammalian orthoreoviruses by possessing a FAST protein and
          protein, expressed in Escherichia coli, generated immunity in   therefore inducing syncytial CPE in cell culture. The evolutionary
          chickens. Wan et al. (2010) showed that the σC DNA vaccine   relationships among avian reoviruses have been widely studied
          orally delivered by attenuated  Salmonella typhimurium elicited   by phylogenetic analysis. Although there are commercially avail-
          antibody production in SPF chicks. Wu et al. (2005) character-  able avian reovirus vaccines for use in commercial poultry, new
          ized the immune response of SPF chickens to orally administered   genetic virus variants continue to emerge making these vaccines
          recombinant  σC protein produced  in  yeast and  recommend   inadequate. Moreover, the increasing diagnosis of avian reovirus
          the development of a transgenic edible plant vaccine for use in   infections in wild birds has raised the possibility of transmission
          poultry. Protein σC of  orthoreovirus  was  expressed in  alfalfa,   between wild birds  and domesticated  poultry. The  application
          Arabidopsis and tobacco plants (Huang et al., 2006; Wu et al.,   of  the  recently  reported  reverse  genetics  system  using  avian
          2009; Lu et al., 2011), but the low expression of antigens in   reoviruses will significantly increase our understanding of avian
          plants limited their practical application in vaccination protocols   reovirus biology and disease.
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