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324  |  Coppo et al.

          Table 11.1  Continued
          Open                                  Predicted   Apparent
          reading                      Predicted   molecular   molecular
          frame                        amino acid  mass   mass
          (ORF)     Protein            length a  (kDa) b  (kDa)    Putative function  Transcribed References
          UL[–1] c  Protein LORF2 (spliced) d  501   57.6  73      Unknown. Nuclear   γ         Ziemann et al. (1998b);
                                       (47 + 454)                  localization       L II      Mahmoudian et al. (2012);
                                                                                                Nadimpalli et al. (2017)
          ICP4      Transcriptional regulator   1491  161.8  200   Gene regulation    IE        Johnson et al. (1995b);
                    Infected cell protein 4                                                     Mahmoudian et al. (2012)
                    (ICP4)
          US10      Virion protein US10  279    30.8     ND        Unknown            E I       Wild et al. (1996)
          sORF 4/3  Protein SORF3      295      32.1     ND        Unknown            L II      Wild et al. (1996);
                                                                                                Mahmoudian et al. (2012)
          US2       Virion protein US2  230     22.3     ND        Unknown – possibly   L I     Mahmoudian et al. (2012)
                                                                   envelope associated
          US3       Serine/threonine protein   477  53.8  ND       Protein phosphorylation   γ1  Kongsuwan et al. (1995);
                    kinase US3                                     – apoptosis – nuclear   E II  Mahmoudian et al. (2012)
                                                                   egress
          UL47 c    Tegument protein   625      67       67.5      Possibly gene regulation  γ2  Kongsuwan et al. (1995);
                    VP13/14 d                                                         E II      Helferich et al. (2007a,b);
                                                                                                Mahmoudian et al. (2012)
          US4 c     Glycoprotein G d   293      32       52        Immune-modulation  γ (α, β)  Kongsuwan et al. (1993a);
                                                                                      L III     Sun and Zhang (2005);
                                                                                                Devlin et al. (2006b,
                                                                                                2010); Helferich et al.
                                                                                                (2007a); Mahmoudian et
                                                                                                al. (2012)
          US5 c     Glycoprotein J d,e  996, 611  107, 67  85, 115,   Virulence. Cell egress  L III  Veits et al. (2003a); Fuchs
                                                         160, 200                     γ2        et al. (2005); Mahmoudian
                                                                                                et al. (2012)
          US6 c     Glycoprotein D d   435      38.7     65, 70    Cell attachment – binds   E/L II  Mahmoudian et al. (2012);
                                                                   to cell surface receptors    Pavlova et al. (2013)
          US7 c     Glycoprotein I d   363      38.2     62        Cell-to-cell spread –   E/L II  Devlin et al. (2006a);
                                                                   complexed with gE to         Mahmoudian et al. (2012);
                                                                   form Fc-receptor             Pavlova et al. (2013)
          US8 c     Glycoprotein E d   500      55.4     75        Cell-to-cell spread –   E/L I  Devlin et al. (2006a);
                                                                   complexed with gI to         Mahmoudian et al. (2012);
                                                                   form Fc-receptor             Pavlova et al. (2013)
          US9 c     Membrane protein US8A d  261  27.9   37, 25                       L III     Mahmoudian et al. (2012);
                                                                                                Pavlova et al. (2013)

          a Based on the sequence of ILTV strain CSW1 (JX646899.1).
          b Where predicted molecular masses have not been published these were estimated using predicted amino acid sequences of the ILTV strain
          CSW1 and a Protein Molecular Weight Calculator tool (http://www.sciencegateway.org/tools/index.html).
          c A gene-specific deletion mutant has been generated for this ORF.
          d Polyclonal antibody has been raised against this protein.
          e Monoclonal antibody has been raised against this protein.
          E, early; E/L, early/late; IE, immediate early; L, late; ND, not determined; P, precursor.

          exhibited an attenuated phenotype, causing less severe clinical   to be undertaken. Also, ultrastructural studies to investigate varia-
          signs than the parent strain and only minor microscopic lesions   tions in viral assembly, maturation and egress associated with the
          in the trachea, despite viral titres in tracheal swabs that were   expression (or lack thereof) of these ILTV unique peptides would
          not significantly different at 6 days after infection (García et al.,   be useful to further characterize their role in these processes.
          2016). Although inconclusive, the data gathered so far indicates   The two other ILTV-specific ORFS, UL0 and UL[–1], located
          that ORFs A–E have important and possibly redundant roles in   upstream to the UL1 ORF, are translated from spliced mRNA
          viral replication and viral egress, and at least for ORF C, its role in   species (Ziemann et al., 1998b). Both pUL0 and pUL[–1] locate
          cell-to-cell spread but not in viral replication per se appears to be   to the nucleus of infected cells (Ziemann et al., 1998b) and are
          associated with the capacity of the virus to cause tissue disruption   categorized as late (γ) transcription genes (Ziemann et al., 1998b;
          and pathology in vivo. Future work to determine whether these   Mahmoudian et al., 2012). Isolation of a deletion mutant lacking
          proteins are only expressed during infection or are a structural   UL0 demonstrated the dispensable nature of this gene, where
          part of the virion (tegument- or membrane-associated) remains   the lack of UL0 resulted in a moderate delay in viral replication
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