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Avian Paramyxoviruses |   107

          APMV-5                                                cell culture, it was observed that all four strains of APMV-6 repli-
          APMV-5 (AMAvV-5) was first isolated in 1974 from an outbreak   cated in a trypsin-independent manner in at least one of the cell
          in budgerigars at Kunitachi, Japan (Nerome et al., 1978). APMV-5   types, but they required trypsin in one or more cell types. This
          is unusual in that it lacks a virion haemagglutinin and does not   suggests the existence of unexpected diversity in the ability of
          grow in the allantoic cavity of embryonated chicken eggs. How-  various cell lines to cleave different F proteins of an APMV (Xiao
          ever, the virus grows in the amniotic cavity of embryonic chicken   et al., 2010).
          eggs and in various cell lines. To date, there are only four reports in   Sequence comparison and cross-HI test results grouped the
          the world on APMV-5 infections (Mustaffa-Babjee, 1974; Gough   APMV-6 strains into two subgroups within a single serotype
          et al., 1993; Yoshida et al., 1997; Hiono et al., 2016). The complete   (Xiao et al., 2010). The strain from red-necked  stint and  the
          genome sequences are available for two of the APMV-5 strains   strain from Italy (IT4524-2) formed one group and the other
          isolated in Japan (Samuel et al., 2010; Hiono et al., 2016). The   strains formed the second group (Xiao et al., 2010; Karamendin
          genome is 17,262 nt long and encodes six genes as other APMVs.   et al., 2013). Recently, sequence analysis of the F gene of 11
          The genome contains a 55-nt leader sequence at the 3′ end and   APMV-6 isolates from wild ducks in Korea also showed exist-
          a 552-nt trailer sequence at the 5′ end. The complete genome   ence of two subgroups within APMV-6 (Choi  et al., 2018).
          sequence of TI strain shares 97% nt sequence identity with the   Another recent study based on the mean inter-populational
          Kunitachi strain (Hiono et al., 2016). The cleavage site of the F   evolutionary distance of the F gene sequence of 24 isolates
          protein  (G-K-R-K-K-RF)  conforms  to the cleavage  site  motif   from China also showed existence of two genotypes (I and II)
          of the ubiquitous cellular protease furin. Consistent with this,   within APMV-6 (Chen et al., 2018). A phylogenetic analysis of
          exogenous protease is not required for virus replication in vitro.   APMV-6 strains is shown in Fig. 3.13.
          Infected of Vero cells by strain Kunitachi showed cell rounding
          and detachment as well as syncytia formation. However, the ICPI   APMV-7
          value of the Kunitachi strain is 0.00, indicating that the virus is   APMV-7 (AMAvV-7) was first isolated from a hunter-killed
          avirulent for chickens, despite containing a multibasic F cleavage   dove (Columba livia) in 1975 in Tennessee, USA (Alexander et
          site.                                                 al., 1981). Since then APMV-7 viruses have been isolated mostly
                                                                from  Columbiformes. Although an  APMV-7 virus was isolated
          APMV-6                                                from a natural outbreak of respiratory disease in commercial
          APMV-6 (AMAvV-6) was first isolated from a domestic duck (A.   turkey breeder flocks in Ohio in 1997, it has not been associated
          platyrhynchos domesticus) in Hong Kong in 1977 (Shortridge et   with severe disease in poultry (Saif et al., 1997). APMV-7 has also
          al., 1980). Since then APMV-6 viruses have been isolated from   been isolated from ostriches (Woolcock et al., 1996). Whether
          a wide range of avian species, including ducks, geese, common   or not the strains of APMV-7 isolated from different avian spe-
          egrets  and red-necked stint worldwide. To  date, complete   cies are the same has not been determined. The genome of the
          genome sequences of eight strains of APMV-6 are available   APMV-7 prototype strain dove/Tennessee/4/75 is 15,480 nt
          (Chang et al., 2001; Xiao et al., 2010; Tian et al., 2012; Kara-  and contains six genes (Xiao et al., 2009). The leader and trailer
          mendin et al., 2013). APMV-6 is different from all other APMV   regions are 55 and 127 nt, respectively. The 3′-leader contains a
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          serotypes in having an SH gene between the F and HN genes   sequence ( AAUUAUUUUUU ) that is identical to the GE
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          (Chang et al., 2001; Xiao et al., 2010). The biological function   signal present at two of the genes. The IGS ranges between 11 and
          of APMV-6 SH gene is not known. However, the SH proteins   70 nt. The F protein has a single basic amino acid at the putative
          of SV5 and mumps virus appear to play a role in blocking TNF-  cleavage site (T-L-P-S-S-RF). The virus grows in only a few
          alpha-mediated apoptosis pathway (Wilson et al., 2006). The   established cell lines, indicating a restricted host range. APMV-7
          SH protein of respiratory syncytial virus (RSV) has ion channel   does not form syncytia or plaques in cell culture and its replica-
          activity (Gan et al., 2008). RSV from which SH gene was deleted   tion does not require exogenous protease. Sequence alignment
          was fully viable in cell culture but was slightly attenuated in mice   and phylogenetic analysis of the predicted amino acid sequence
          and chimpanzees (Bukreyev et al., 1997; Whitehead et al., 1999).   of APMV-7 proteins with the cognate proteins of the viruses of
          APMV-6 strain duck/Hong Kong/18/199/1977 (Hong Kong)   other members of subfamily Avulavirinae showed that APMV-7 is
          is the prototype strain.                              more closely related to APMV-2, -5, -6, -8, -10, -14, -15, and -20
            Six out of the eight available APMV-6 complete genome   than to other APMV serotypes.
          sequences, including strain Hong Kong, contain a genome of
          16,236 nt. However, two sequences, one from a stint in Japan and   APMV-8
          the other from a duck in Italy, have 16,230 nt. The six-nucleotide   APMV-8 (AMAvV-8) was first isolated from a Canadian goose
          difference between the two viruses is due to a deletion in the non-  (Branta  canadensis) in 1976 in Delaware, USA (Cloud and
          coding region of the F gene (Xiao et al., 2010).      Rosenberger, 1980). Another APMV-8 strain was isolated from
            In  cell  culture,  APMV-6  induced  cell  rounding  and  detach-  a feral pintail duck (A. acuta) in 1978 in Wakuya, Japan (Yamane
          ment, and not syncytia formation (Xiao et al., 2010). The F   et al., 1982). APMV-8 has not been isolated from domestic poul-
          protein cleavage site of the Italian strain IT4524-2 has dibasic aa   try. Determination of the first complete genome sequence of
          (R-E-P-RL), compared with the monobasic F cleavage site of   APMV-8 strain goose/Delaware/1053/76 (prototype strain) was
          P-E-P-RL in other strains. Among four APMV-6 strains tested in   independently published by two research groups (Mueller et al.,
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