ROUTES OF EXCRETION IN FLYING-FOXES
Urine was confirmed as the major route of HeV excretion from flying-foxes through an extensive analysis of nearly 3000 individuals (based on both the rate of HeV detection and amount of viral RNA across a range of sample types; Edson et al.
2015b). Following urine, the highest levels of detection were observed in serum and packed haemocytes, then faecal, nasal and oral swabs. In addition to evidence for haematogenous spread, the predilection of HeV for the spleen in a retrospective study of archived bat tissues suggests this tissue may play an important role in naturally acquired HeV infection in flying-foxes (Gold- spink et al. 2015). A lack of viral RNA detection in the foetal tissues of four HeV-positive females in that study contrasted earlier studies (Halpin et al. 2000; Williamson et al. 2000) and suggests that vertical transmission is not a consistent mode of transmission in naturally infected flying-foxes. Multiple studies have suggested that the direct contamination of horses (via ocular, nasal and buccal mucous membranes) under trees in which flyingfoxes are feeding is a plausible alternative mode of transmission to indirect contact of infectious materials via ingestion of contaminated pasture (Edson et al. 2015b; Martin et al. 2015). There is no evidence that bat flies (Nycteribiid ectoparasites) play a role in HeV transmission (Vidgen et al. 2017).4.
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