Japanese encephalitis virus (JEV) (Flavivirus: Flaviviridae) is a leading cause of encephalitis in eastern and southern Asia. The virus is maintained in a zoonotic cycle between ardeid wading birds and/or pigs and Culex mosquitoes. The primary mosquito vector of JEV is Culex tritaeniorhynchus, although species such as Cx. gelidus, Cx. fuscocephala, and Cx. annulirostris are important secondary or regional vectors. Control of JEV is achieved through human and/or swine vaccination, changes in animal husbandry, mosquito control, or a combination of these strategies. This review outlines the ecology of JEV and examines the recent expansion of its geographical range, before assessing its ability to emerge in new regions, using the hypothetical establishment in the United States as a case study.

Ecology and Geographical Expansion of Japanese Encephalitis Virus

First discovered in 1947, Zika virus (ZIKV) infection remained a little-known tropical disease until 2015, when its apparent association with a considerable increase in the incidence of microcephaly in Brazil raised alarms worldwide. There is limited information on the key factors that determine the extent of the global threat from ZIKV infection and resulting complications. Here, we review what is known about the epidemiology, natural history, and public health effects of ZIKV infection, the empirical basis for this knowledge, and the critical knowledge gaps that need to be filled.

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Assessing the global threat from Zika virus

Emerging arboviruses: Why today?

Since April 2010, a severe outbreak of duck viral infection, with egg drop, feed uptake decline and ovary-oviduct disease, has spread around the major duck-producing regions in China. A new virus, named BYD virus, was isolated in different areas, and a similar disease was reproduced in healthy egg-producing ducks, infecting with the isolated virus. The virus was re-isolated from the affected ducks and replicated well in primary duck embryo fibroblasts and Vero cells, causing the cytopathic effect. The virus was identified as an enveloped positive-stranded RNA virus with a size of approximately 55 nm in diameter. Genomic sequencing of the isolated virus revealed that it is closely related to Tembusu virus (a mosquito-borne Ntaya group flavivirus), with 87–91% nucleotide identity of the partial E (envelope) proteins to that of Tembusu virus and 72% of the entire genome coding sequence with Bagaza virus, the most closely related flavivirus with an entirely sequenced genome. Collectively our systematic studies fulfill Koch's postulates, and therefore, the causative agent of the duck egg drop syndrome occurring in China is a new flavivirus. Flavivirus is an emerging and re-emerging zoonotic pathogen and BYD virus that causes severe egg-drop, could be disastrous for the duck industry. More importantly its public health concerns should also be evaluated, and its epidemiology should be closely watched due to the zoonotic nature of flaviviruses.

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Duck egg-drop syndrome caused by BYD virus, a new Tembusu-related flavivirus.

Among animal viruses, arboviruses are unique in that they depend on arthropod vectors for transmission. Field research and laboratory investigations related to the three components of this unique mode of transmission, virus, vector, and vertebrate host, have produced an enormous amount of valuable information that may be found in numerous publications. However, despite many reviews on specific viruses, diseases, or interests, a systematic approach to organizing the available information on all facets of biological transmission and then to interpret it in the context of the evolutionary process has not been attempted before. Such an attempt in this review clearly demonstrates tremendous progress made worldwide to characterize the viruses, to comprehend disease transmission and pathogenesis, and to understand the biology of vectors and their role in transmission. The rapid progress in molecular biologic techniques also helped resolve many virologic puzzles and yielded highly valuable data hitherto unavailable, such as characterization of virus receptors, the genetic basis of vertebrate resistance to viral infection, and phylogenetic evidence of the history of host range shifts in arboviruses. However, glaring gaps in knowledge of many critical subjects, such as the mechanism of viral persistence and the existence of vertebrate reservoirs, are still evident. Furthermore, with the accumulated data, new questions were raised, such as evolutionary directions of virus virulence and of host range. Although many fundamental questions on the evolution of this unique mode of transmission remained unresolved in the absence of a fossil record, available observations for arboviruses and the information derived from studies in other fields of the biological sciences suggested convergent evolution as a plausible process. Overall, discussion of the diverse range of theories proposed and observations made by many investigators was found to be highly valuable for sorting out the possible mechanism(s) of the emergence of arboviral diseases.

Biological Transmission of Arboviruses: Reexamination of and New Insights into Components, Mechanisms, and Unique Traits as Well as Their Evolutionary Trends

The possible role of pigs as arbovirus maintenance hosts and their importance as amplifier hosts was studied. Blood samples from 464 pigs of all ages collected in 1962 and 1964 were tested against 10 arboviruses. Antibodies to Japanese encephalitis and Getah viruses were particularly prevalent and their calculated monthly infection rates were 19-5% and 13-3% respectively. In 1969, 447 pigs were bled monthly throughout the year and the infection rates for Japanese encephalitis virus were calculated in pigs during the first year of life. Infection rates were not uniform throughout the year; the rate increases as the pig grew older and there was a marked seasonal increase in the infection rate in the period from November to January. This coincided with the seasonal major population peak of Culex tritaeniorhynchus following intense breeding of this mosquito prior to rice planting. It is suggested that, in Sarawak, the pig acts as a maintenance host of Japanese encephalitis in a cycle involving C. gelidus mosquitoes and also acts as an important amplifier host towards the end of the year in a cycle involving C. tritaeniorhynchus. It is further suggested that Getah virus is maintained in a similar cycle between C. tritaeniorhynchus and pigs.

Arbovirus infections in Sarawak: the role of the domestic pig

1,844 of 4,109 sera collected from children in 26 schools on the Kano Plain, Nyanzo Province, Kenya during May and June 1969 were tested for antibodies to a variety of arboviruses. Over 60% of the children had HI antibodies against both Chikungunya and O'nyong-nyong viruses and 45% had antibody levels of 1 : 160 or greater against O'nyong-nyong suggesting recent infection with this virus. 35% of children aged less than 6 years had O'nyong-nyong virus antibodies and O'nyong-nyong was probably the virus responsible for most of the group A virus antibody. The average annual probability of infection with O'nyong-nyong virus was calculated at 9·6% and the virus is probably endemic in the area. Very few sera were found with group B virus antibodies but West Nile and probably Zika virus is thought to be endemic at a low level. Bunyarnwera group viruses are active in the area and Bwamba virus also appears to be endemic. Nyando virus antibodies were found in a considerable number of children aged 10 years or older and an epidemic probably occurred some 10–15 years previously; but there is no evidence that transmission is active at present.

Large scale irrigation and arbovirus epidemiology, Kano plain, Kenya II. Preliminary serological survey

449 human sera collected in a Land Dyak village were tested for antibodies to 11 arboviruses. Japanese encephalitis and dengue virus antibodies were particularly prevalent. The rates of infection with these viruses were estimated to be 5-2% per annum for Japanese encephalitis, 8-8% for dengue 1 and 4-3% for dengue 2. Chikungunya virus antibodies were quite common with an annual infection rate of the order of 5% per annum. Infections with other Group A and B and Bunyamwera group viruses were generally at a low level.

Janice Day

Papers

Arbovirus infections in Sarawak: the role of the domestic pig

Large scale irrigation and arbovirus epidemiology, Kano plain, Kenya II. Preliminary serological survey

Arbovirus infections in Sarawak, October 1968–February 1970: Human serological studies in a Land Dyak Village