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Avian Adenovirus |   307

          Prevention and control                                against egg-drop syndrome with good results. When the birds
          Fowl adenoviruses are associated with mild and severe diseases   are vaccinated at 14–16 week of age, the immunity will last up
          in poultry; IBH, HPS and GE are economically important   to a year (Baxendale et al., 1980; Kaleta et al., 1980; Solyom et
          (McFerran and Smyth, 2000). EDSV is widely distributed in   al., 1982; Cook, 1983). EDSV fibre protein expressed by E.coli
          domestic and wild ducks and geese. Infection of chickens with   induced a high neutralizing antibody titre after two vaccinations
          EDSV causes major economic losses due to its direct effect on   that lasted more than 50 weeks of lay (Gutter et  al., 2008). A
          egg production and eggshell quality (Kang et al., 2017). HEV is   further engineered fibre protein expressed in  E. coli prevented
          associated with acute haemorrhagic gastro-intestinal disease of   clinical signs of the disease when the vaccinated chickens were
          6–11-week-old turkeys and ablation of the humoral and cellular   challenged with a pathogenic EDSV (Harakuni et al., 2016).
          immune response (Dhama et al., 2017).                   There are live attenuated virus vaccines available that would
            In general, avian adenoviruses are quite resistant to inactivation   provide good protection against HE of turkeys (Pierson and
          and persist in the environment for a long period of time. Manage-  Fitzgerald, 2013; Dhama et al., 2017). The orally administered
          ment practices and good biosecurity are very important, however   cell culture vaccine is commercially available and recommended
          because of the vertical viral transmission and co-infection with   in many countries (Fadly et al., 1985; Barbour et al., 1993).
          other viruses, such as IBDV and CAV, which could enhance path-  Genetic engineering techniques have been also utilized to gen-
          ogenicity (Hess, 2013), the control of the diseases often seems to   erate subunit vaccines against HEV. The earlier ones expressed
          be difficult. Therefore, vaccination is generally recommended and   the major immunogenic protein, the hexon, either by a bacterial
          practised with promising successes.                   system or through a fowlpox virus vector (van den Hurk and van
            Protective antibody response is detectable after one week of   Drunen Littel-van den Hurk, 1993; Cardona et al., 1999, 2001).
          infection with fowl adenoviruses (Hess, 2013). The first suc-  A recombinant penton-fibre knob subunit vaccine has also been
          cessful protection of chickens against inclusion body hepatitis   developed and it is marketed in certain countries (Pitcovski et al.,
          through vaccination was reported over 40 years ago (Fadly and   2005).
          Winterfield, 1975). Both live and inactivated vaccines containing   Commercial vaccines, inactivated or live attenuated, are avail-
          different serotypes have been evaluated. In Australia, an FAdV-8b   able to control diseases caused by avian adenoviruses. Inactivated
          strain-based commercially available live vaccine is given to broiler   EDSV vaccines are available in many countries including the
          breeders. However, IBH outbreaks in broilers were reported and   USA, as either monovalent or multivalent (e.g. Agrovet, Volvac   ®
          it was shown that the disease resulted from alternative FAdV   AC, Boehringer Ingelheim Vetmedica Inc, etc.). Other vaccines
          strains of different species (Steer et al., 2011). Inactivated, wild   to EDS, available in other countries but not in the USA, are
                                                                                                     ®
          type FAdV-8 and FAdV-11 strains have been successfully used   prepared as multivalent vaccines (e.g. Volvac ) for infectious
          to confer protection via maternal antibodies to broiler-breeders   bronchitis and HPS/IBH (FAdV-4/8). Subunit vaccines for EDS
          when grandparents were vaccinated (Alvarado et al., 2007). In   (knob fibre protein), delivered by subcutaneous or intramuscular
          some countries, for example Canada, the broiler breeders are vac-  injections, have also been licensed in many countries including
          cinated with autogenous vaccines against IBH, resulting in greatly   the USA. Such vaccines are also prepared as multivalent ones for
          reduced mortality (Brash, 2012). In many countries, vaccines   other poultry diseases namely Newcastle disease and infectious
          derived from liver homogenate are commonly used against HPS/  bronchitis (e.g. Phibro Vaccines).
          IBH (Asthana et al., 2013). Effective protection of the progeny   Vaccines to HE can be generated from spleen homogenates
          of chickens against HPS/IBH is achieved by dual vaccination   obtained from 6-week-old turkeys inoculated with an avirulent
          of breeders with killed FAdV-4 and CAV (Toro et al., 2002).   HEV strain. However, live-attenuated vaccines obtained from
          Virulent FAdV-4 isolates were attenuated by adapting to grow in   cell culture systems or originating from other avian species are
          a quail fibroblast cell line (QT-35) or in SPF chicken embryos   available commercially (Dhama et al., 2017). For example, some
          and applied as vaccines (Schonewille et al., 2008; Mansoor et al.,   HE vaccines are of pheasant origin (e.g. Ceva Adenomune™) or
          2011). Immunization resulted in reduced mortality upon FAdV-4   propagated in Marek’s disease induced B-lymphoblastoid cell
          challenge of the chicks but did not prevent virus shedding. Simi-  line (MDTC-RP19) of turkey origin (e.g. Arko Laboratories
          larly, administration of FAdV-1 to SPF broiler chickens did not   Ltd). Both vaccines are administered in the drinking water to
          prevent virus shedding but protected against clinical signs of GE   30-day-old turkeys. Subunit vaccines that consist of the fibre
          (Grafl et al., 2014). In addition to whole virus vaccines, subunit   knob protein have also been developed to control HE (e.g. Abic
          vaccines with the primary FAdV antigens, the hexon, penton base   Biological Laboratories Teva Ltd).
          and fibre were also reported. When baculovirus expressed puri-  Autogenous vaccines are formalin-inactivated liver homoge-
          fied fibre 2 protein of FAdV-4 was administered intramuscularly   nates used to control mainly HPS (Mansoor et al., 2011).
          to 1-day-old chickens the mortality was reduced to 3.5% after   Commercial inactivated vaccines to control HPS/IBH are avail-
          challenge with a virulent virus (Schachner et al., 2014). Similarly,   able in many countries but not in the USA. These vaccines are
          E. coli expressed purified FAdV-4 penton base provided 90% pro-  directed mainly to FAdV-4 and FAdV-8a and available as either
          tection against subcutaneous challenge 2 weeks later (Shah et al.,   monovalent or bivalent versions. These vaccines are also available
          2012).                                                as multivalent forms for the control of other poultry diseases (e.g.
            Oil adjuvanted killed vaccines are used for the immunization   Avimex  Animal  Health  and  Boehringer  Ingelheim  Vetmedica
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