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134  |  Liu et al.

          identified in Europe, perhaps the ones of major importance are   ribosomal frame-shifting, (c) the structural and accessory genes
          4/91,  793B,  CR88  (Gough et al.,  1992;  Parsons et al.,  1992).   downstream of the replicase gene are expressed via a 3′-nested
          Some IBV variants may be geographically restricted, and multi-  subgenomic messenger RNAs (sgRNAs) and (d) a collection
          ple studies have employed molecular methods to identify IBV   of viral enzymatic activities are encoded within the replicase-
          variants unique to the region (Escorcia et al., 2000; Collison et al.,   transcriptase protein products.
          2001; Gelb et al., 2001; Alvarado et al., 2005).         The classification systems of IBV strains are divided into two
                                                                major  groups, namely  the functional and  non-functional  tests
                                                                (Table 5.1). Functional tests look into the biological functions
          Nomenclature                                          of the said IBV strains, and categorize IBV into various patho-
          IBV is an enveloped, positive sense, single-stranded RNA virus   types, protectotypes and antigenic types. On the other hand, the
          classified under the family Coronaviridae of the order Nidovirales,   non-functional tests probe into the viral genome, and this usu-
          along with families Arteriviridae, Mesoniviridae, and Roniviridae   ally results in grouping IBVs based on genotype (de Wit, 2000;
          (International Committee on Taxonomy of viruses, http://www.  Valastro et al., 2016).
          ictvonline.org/virustaxonomy.asp). Virus taxonomy of the family   Sequence comparisons of the complete genome, spike (S)
          Coronaviridae is shown in Fig. 5.1. Nidoviruses are set apart   protein subunits 1 and 2 (S1 and S2), envelope (E) protein, mem-
          from other RNA viruses by four distinctive characteristics: (a)   brane (M) protein and nucleocapsid (N) protein of IBV strains
          an invariant genome organization comprising a large replicase   from distinct geographical regions have contributed great strides
          gene occupying the 5′ two-thirds of the viral genome, (b) transla-  in the construction of the IBV phylogenetic tree (Fig. 5.2) (Lin
          tion of the replicase-transcriptase polyprotein is performed via   et al., 2016).



                                                                    Nidovirales

                                                                   Coronaviridae



                                                Coronavirinae                            Torovirinae






                       Alphacoronavirus Betacoronavirus  Gammacoronavirus  Deltacoronavirus  Torovirus  Bafinivirus




                                                 Infectious bronchitis virus (IBV)

          Figure 5.1  Taxonomy of the family Coronaviridae. Infectious bronchitis virus (IBV) is a Gammacoronavirus, under the family Coronaviridae
          within the order Nidovirales.

                            Figure 1. Taxonomy of the family Coronaviridae. Infectious bronchitis virus (IBV)
          Table 5.1  Classification of infectious bronchitis virus (IBV) into pathotypes, protectotypes, antigenic types and genotypes
                            is a Gammacoronavirus, under the family Coronaviridae within the order Nidovirales.
                            Parameter measured                         Features
          Functional tests
          Pathotypes        Clinical signs, gross lesions, and virus pathogenicity;   Pros: practical in the field for vaccine strategy
                            same pathotype when both tested strains induce similar
                            pathological signs
          Protectotypes     Complete immune response against an IBV strain; same   Pros: provides valuable information about vaccine efficacy
                            protectotype when strains induce protection against each   Cons: laborious and expensive; requires high-level facilities
                            other                                      for vaccination-challenge studies
          Antigenic types   Reaction between IBV strain and chicken-induced IBV   Cons: less practical when more IB variants are found in the
          (serotypes and    serotype-specific antibodies; same serotype when   area as every serotype requires its own neutralization test.
          epitope type)     heterologous neutralization titres differ less than 20-fold   For new IB strains, an antiserum has to be raised in SPF
                            from homologous titres                     birds.
          Non-functional tests
          Genotypes         Genetic characterization of viral genome; same genotype   Pros: objective; provides useful information for
                            when sequence of tested strain matches     epidemiological studies
          IB, infectious bronchitis; IBV, infectious bronchitis virus; SPF, specific pathogen free.
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