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Chapter 3: Transmission / Natural cycle


          Vertical TBEV transmission in               Outbreaks   due   to   alimentary   virus
          vertebrates                                 transmission are known from Eastern, Central
                                                      and  Southern  Europe, 122,123   and  have  to  be
                                                      considered  particularly  in  cases  of  local
          TBEV  transmission  from  mother  to  her   epidemics. 123–125
          offspring in small rodents, e.g., red voles (M.
          rutilus),  was  shown  for  naturally  infected
          reservoir  hosts  as  well  as  after  experimental   In  summary,  all  these  ways  of  transmission
                                                      together  ensure  the  long-term  perpetuation
          infection with different sublethal doses of the
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          virus.  TBEV  RNA was detected in up to 90%   of TBEV in many foci.
          of  the  newborn  rodents,  240–280  days  after
          experimental  infection  of  their  parents,  by   Changes  in  environmental  conditions,  e.g.
          real-time polymerase chain reaction (RT-PCR),   temperature,  strengthen  the  importance  of
          enzyme-linked  immunosorbent  assay  (ELISA),   one  possibility  or  of  another.  Thus,  the
          and  bioassays.  The  small  amounts  of  TBEV   development  of  eggs  in  infected  females
          RNA  detected  in  the  embryos,  placenta,  and   under high temperature conditions leads to an
          blood  serve  as  evidence  of  prenatal  trans-  increase  in  the  proportion  of  transovarially
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          mission. Postnatal transfer of the virus might   infected  larvae.    An  increase  in  summer
          occur through the rodent’s milk. Vertical virus   temperature  increases  the  frequency  of
          transmission may occur before, during, and/or   occurrence  of  engorged  vector  tick  females
          after  birth  of  the  baby  rodents  with  a  high   with traces of male bites, with the occurrence
          frequency.  In  natural  foci,  this  could  ensure   of  bites  being  higher  among  southern
          long-term  persistence  of  TBEV  in  mammal   populations of I. persulcatus (up to 10% in the
          hosts  without  involving  any  arthropod   region  of  Zhiguli)  than  among  northern  ones
                 119
          vectors.                                    (less  than  2%  in  the  Krasnoyarsk  popular-
                                                          118
                                                      tion).   Therefore,  under  changing  abiotic
                                                      conditions, different homeostatic mechanisms
          Reservoir  incompetent  hosts  do  not  directly
          participate in virus transmission, but can play   become  apparent.  Tick  behavior  can  also  be
                                                      an important factor affecting the effectiveness
          an  important  role  in  the  maintenance  of
                                                      of TBEV transmission, with the TBEV infection
          natural  foci.  The  density  of  reservoir-
          incompetent hosts may have either a positive   in the tick itself  possibly serving to stimulate
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          effect on virus transmission, by amplifying the   the questing activity of the tick.
          tick  population,  or  a  negative  (‘dilution’)
          effect, as tick bites on a reservoir-incompetent
          host cannot lead to virus transmission. 98,120
                                                      The natural cycle
          Alimentary route of transmission            The  natural  cycle  of  TBEV  is  highly  complex,
                                                      and  many  details  remain  obscure.  The  three
                                                      prevailing  TBEV  subtypes  overlap  in  some
          Humans  mostly  become  infected  with  TBEV   areas,  they  all  have  multiple  mammalian
          via  tick  bites,  but  viral  transmission  is  also   reservoir hosts and various tick vectors, and in
          possible via the consumption of unpasteurized   some  areas  these  subtypes  occur  sym-
                               43
          goat, cow and sheep milk.  Approximately 1%
                                                      patrically.  Humans  are  not  included  in  these
          of all TBEV infections in humans are probably   natural  cycles,  but  may  enter  those  trans-
          acquired by consuming infected unpasteurized   mission cycles inadvertently.
          milk and milk products from infected livestock,
          particularly goats. 121                     Small mammals and vector ticks play a central
                                                      role in the natural cycle of TBEV, but birds and


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