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Marek’s Disease Virus |   349
                                                                     DNA Pol

                                                          UL
                     TRL                                                                            IRL
                           UL2 UL5  UL8 UL9  UL12    UL23   UL29 UL30  UL39 UL40 UL42  UL50 UL52
                                                                          RR



                                                            DNA H-P

          Figure 12.1  DNA replication associated genes of the MDV. The unique long (UL) region is flanked by terminal (TRL) and internal repeat
          long (IRL) regions. The location of the origins of replication is indicated by vertical arrows. The essential and non-essential DNA replication
          associated genes are listed (black and grey, respectively). The subunits of DNA polymerase (DNA Pol), DNA helicase and primase (DNA H-P),
          and ribonucleotide reductase (RR) are shown.
                       Figure 1

          Table 12.4  MDV proteins involved in replication.
                   HSV                               Alternate     Essential for DNA
          MDV gene  homologue  Protein               abbreviation  synthesis     Major function
          MDV021   UL9        Origin-binding protein  OBP          Yes           Origin-binding protein
                                                                                 3’–5′ DNA helicase
          MDV042   UL29       Major single-stranded   ICP8, SSB,   Yes           Single-stranded DNA-binding protein;
                              DNA-binding protein    MDBP                        Single-strand annealing protein;
                                                                                 Stimulates DNA polymerase,
                                                                                 helicase-primase, and origin binding protein
          MDV017   UL5        DNA helicase-primase                 Yes           Subunit of helicase-primase;
                                                                                 Contains helicase motifs
          MDV020   UL8        DNA helicase-primase                 Yes           Subunit of helicase-primase;
                                                                                 Stimulates primer synthesis, interacts with
                                                                                 ICP8 and OBP proteins
          MDV066   UL52       DNA helicase-primase                 Yes           Subunit of helicase-primase;
                                                                                 Contains primase motifs
          MDV043   UL30       DNA polymerase         Pol           Yes           Catalytic subunit of DNA polymerase;
                                                                                 3’–5- exonuclease, RNase H
          MDV055   UL42       DNA polymerase                       Yes           Processivity subunit of DNA polymerase;
                                                                                 Double-strand DNA-binding protein
          MDV024   UL12       Alkaline nuclease      5′–3′ Exo     No            Endo- and exonuclease;
                                                                                 Interact with ICP8;
                                                                                 Viral recombinase
          MDV014   UL2        Uracil DNA glycosylase  UDG          No            Uracil-DNA glycosylase
          MDV063   UL50       Deoxyuridine triphosphatase  dUTPase  No           Deoxyuridine triphosphatase
          MDV036   UL23       Thymidine kinase       TK            No            Non-specific nucleoside kinase
          MDV052   UL39       Ribonucleotide reductase  RR1, ICP6  No            Large subunit of ribonucleotide reductase
          MDV053   UL40       Ribonucleotide reductase  RR2        No            Small subunit of ribonucleotide reductase




          Two-subunit DNA polymerase (MDV043/UL30 and           Proteins associated with viral DNA replication
          MDV055/UL42, Pol)                                     Although these proteins are not required for DNA replication in
          MDV DNA polymerase exists as a heterodimer, which includes   cultured cells, they are involved in nucleotide metabolism and
          the protein products of MDV043 or UL30 and MDV055 or UL42   have certain effect on the cell growth and viral spread.
          genes. UL30, a 1180 amino acid long protein, possesses both 3′–5′
          exonuclease and RNase H activities. On the other hand, UL42, a   Alkaline nuclease (MDV024/UL12)
          488 amino acid long phosphoprotein, possesses double-strand   MDV alkaline nuclease is encoded by MDV024, a homologue of
          DNA binding activity, and works as the DNA polymerase proces-  HSV-1 UL12 (Osterrieder, 1999). It has endo- and exonuclease
          sivity factor (Sui et al., 1995).                     activities and shows no sequence specificity (Banks et al., 1983).
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