Rodents have long been notorious for spreading disease among humans. Often the animals can carry viruses and transmit them to humans without becoming ill. Certain species thrive in cities and agricultural areas where they come in close contact with humans; this creates many opportunities to spread infection. As humans urbanize and farm larger swaths of previously wild lands, the risk of rodent-transmitted infections increases. As a result, some scientists are working to identify viruses carried by rodents in human settlements and hopefully prevent them from spreading to humans.

The mammarenavirsuses are a group of rodent-transmitted viruses that commonly cause illness in people in Africa and Latin America. Each year, one such virus – the Lassa virus –sickens as many as 300,000 people in Africa and kills 5,000. So far, only two mammarenaviruses have been found in Asia: one called the Wēnzhōu virus and another called LCMV. However only LCMV is known to cause human illness and many cases of illness caused by mammarenaviruses in Asia may go undetected because they often cause mild symptoms similar to the common cold.

Blasdell et al. have now tested lung samples from 20 species of rodents collected at 7 sites in Cambodia, Thailand, and Laos to look for molecules produced by mammarenaviruses. The tests revealed a strain of Wēnzhōu virus circulating in Cambodian rats that often live in urban areas. A new mammarenavirus was also detected in rodents that live in Thai rice fields. However, infecting wild and domestic rodents with the viruses in the laboratory did not cause many noticeable signs of illness.

Blasdell et al. then tested samples from Cambodian patients who either had influenza-like symptoms or more serious symptoms that are associated with a condition called Dengue fever (which is common in the area). Some patients with respiratory symptoms tested positive for the Wēnzhōu virus. Because the symptoms are mild and similar to those of other common diseases it is likely that the Wēnzhōu virus may be spreading more widely among humans in Asia.

The next challenges are to provide a better estimate of the frequency of this disease in the human population in Asia and to describe the full spectrum of disease that might be associated with this newly discovered infectious disease.

Rodents of several species are known hosts of numerous zoonotic pathogens (Luis et al., 2013), and are also amongst the peridomestic ensemble that benefit from how humans are modifying the landscape (Shochat et al., 2006). Their increased presence will amplify human-rodent encounter rates and opportunities for zoonotic transmission, likely creating an increased risk for human health (Young et al., 2014). A concerted effort to establish which human pathogens are present in the environments heavily utilized by humans would therefore be beneficial for the prevention, control and where possible, treatment of such zoonoses.

Mammarenaviruses are predominantly rodent-borne viruses, several of which have been associated with human disease. In Western Africa, Lassa virus (LASV) infection causes an estimated 100,000 to 300,000 infections and approximately 5000 deaths annually (McCormick et al., 1987), whilst Junín virus (JUNV) causes regular seasonal outbreaks of viral haemorrhagic fever in Argentina (Gómez et al., 2011). So far only Wēnzhōu virus and lymphocytic choriomeningitis virus (LCMV) have been conclusively demonstrated to be present in Asia (Li et al., 2014; Morita et al., 1996; 1991). Wēnzhōu virus was identified in China in rodents belonging to four Rattus species, Niviventer niviventer and in Asian house shrews (Suncus murinus), but has not yet been associated with disease. However, LCMV is zoonotic, is primarily hosted by the widespread house mouse (Mus musculus), and consequently has the widest known distribution of any mammarenavirus (Buchmeier et al., 2007). Human infection with LCMV predominantly results in a relatively mild influenza-like illness (Bonthius, 2012; Macneil et al., 2012; Centers for Disease Control and Prevention (CDC), 2012). This propensity of some mammarenaviruses to cause mild disease with symptoms common to those found for many other viral infections, illustrates the potential for these infections to go undiagnosed or misdiagnosed. That human infections caused by arenaviruses other than LCMV have not previously been detected in Southeastern Asia (Charrel and de Lamballerie, 2003) should therefore not be considered as evidence of absence. As each mammarenavirus is primarily adapted to rodents of distinct species or to closely related rodents of diverse species (Gonzalez et al., 2007), the apparent absence of these viruses from a region with high rodent biodiversity (Pagès et al., 2010) is surprising. A large proportion of infectious diseases remain undiagnosed in resource poor countries where people rarely visit healthcare professionals and where clinicians often do not have the training or laboratory support for accurate diagnosis (Mueller et al., 2014). Here, in the course of a survey for rodent-borne mammarenaviruses, we identified a genetic variant of the recently identified Wēnzhōu virus in Cambodia - provisionally named Cardamones - in rodents belonging to two widespread species, brown rats (Rattus norvegicus) and Pacific rats (R. exulans), which are commonly found in proximity to humans. We also identified a novel mammarenavirus species in Thailand in rice-field associated greater bandicoot rats (Bandicota indica), Savile’s bandicoot rats (B. savilei) and an Indomalayan niviventer (Niviventer fulvescens). In addition, we detected Wēnzhōu virus in several Cambodian patients presenting with fever and respiratory symptoms, indicating that this virus may be causally related with human disease.

The identification of a novel mammarenavirus in Thai rodents and of a genetic variant of Wēnzhōu virus in Cambodian rodents is the first unequivocal evidence of mammarenaviruses in Southeastern Asia. The identification of the latter virus in Cambodia in individuals of R. exulans sympatric to infected individuals of the known host, R. norvegicus, expands both the geographic and host range of this mammarenavirus species. Although LCMV was not identified in this study (probably due to the absence of its host the house mouse from large regions of Southeastern Asia (Aplin et al., 2003) and the study sites in particular) it is also proposed to have an Asian origin (Albariño et al., 2010). Despite this shared geographic heritage, both Wēnzhōu virus and LORV appear to be more closely related to the African LASV/IPPYV group viruses. This is also surprising from the viewpoint of the rodent-mammarenavirus co-evolution hypothesis. Based on this hypothesis a closer relationship would be expected between LASV/IPPYV and LCMV, since the rodent hosts (Arvicanthis, Mastomys and Praomys species) (Childs et al., 1993; Lecompte et al., 2005) of the African mammarenaviruses are more closely related to Mus spp. than they are to the Asian Rattini tribe (Lecompte et al., 2008). As such this finding alongside the recent discovery of divergent arenaviruses in boid snakes (recently classified as reptarenaviruses, the sister genus to mammarenaviruses) and more in depth analyses of rodent-mammarenavirus relationships, suggest that the history of the family Arenaviridae is more complex and the theory of co-evolution less robust than initially thought (Coulibaly-N'Golo et al., 2011; Irwin et al., 2012; Stenglein et al., 2012).

Although the host status of rodents of several Rattus spp. could be called into question based on the pathologies observed in some animals experimentally infected with Cardamones variant of Wēnzhōu virus, detrimental effects of infection have been observed in the natural host of another mammarenavirus, Mopeia virus (Borremans et al., 2011), and in the current study may have resulted from the unnatural route of infection. In addition, the successful experimental infection and transmission of Wēnzhōu virus in rodents of two of the proposed host species, alongside the detection of this virus in rodents of several Rattus spp. from two geographically remote locations (Li et al., 2014), support their status as the reservoir hosts. Due to the relative host specificity of mammarenaviruses, confirmation of the host species of each of these viruses has important implications for their geographic distributions. Many of the species in which Wēnzhōu virus or LORV have been detected are widely distributed (Aplin et al., 2003; Ruedi et al., 1996; Wodzicki and Taylor, 1984). Rodents of these species are also associated with either agriculture (Bandicota spp.) or urban settings (Rattus spp.) (Aplin et al., 2003) and as such are likely to be among the few species that may benefit from urbanisation and agricultural intensification. This could ultimately result in increases in both the distribution of their viruses and in their opportunities for zoonotic transmission.

Our results suggest that the ecology of both viruses, through their association with peridomestic rodents of several species, mirrors that observed for known zoonotic mammarenaviruses. Like LORV, JUNV is associated with agricultural pest species (Busch et al., 2000), whilst Wēnzhōu virus, LASV and LCMV are all hosted by rodents with a preference for human dwellings (McCormick et al., 1987; Pocock et al., 2004). This association with highly utilised human habitats indicates that humans are potentially exposed to both viruses on a regular basis as suggested by the relatively high seroprevalence observed in both the patient and the control group. Therefore, the identification of human Wēnzhōu virus infection is perhaps unsurprising from an epidemiological viewpoint, but nevertheless, represents the first detection of human mammarenavirus infection in Asia. The results from this study suggest that human infection with Wēnzhōu virus, or a closely related virus, occurs widely throughout Cambodia, and may possibly result in disease in some cases.

Confirmation of causality however is made problematic by the mild and indistinctive clinical presentation observed, the detection of co-infecting pathogens capable of producing a similar clinical picture in three out of six cases (although rhinovirus infection is not always symptomatic) (Buecher et al., 2010) and the limited number of cases. Although virus was never detected in healthy individuals and all patients with confirmed Wēnzhōu virus infections experienced similar symptoms of acute febrile bronchitis, the non-detection of co-infecting agents in three cases despite extensive testing does not mean that other untested or unknown pathogens were not present. However the clinical picture observed was similar to that reported for milder LCMV infection (Baum et al., 1966; Knust et al., 2014). If like LCMV, Wēnzhōu virus rarely causes distinctive severe disease, then despite the high apparent incidence of the infection indicated by the retrospective serological study, causality is unlikely to be confirmed without further surveys.

Although our findings indicate that novel mammarenaviruses are present in Southeastern Asia and that humans appear to be regularly infected in the region, there are several issues that still need to be addressed. A major one was our lack of success in isolating both Cardamones variant of Wēnzhōu virus and LORV in tissue culture. This was surprising as samples clearly contained viable virus as evidenced by successful experimental infection of rats. It was also inhibitive to downstream analyses, as a successful method of isolation could greatly aid in the development of serological tests and in confirming proof of causality in suspected cases of human disease. The reasons behind our inability to isolate these viruses are unknown. It is possible that the viruses may have replicated too slowly to be detected by the methods used but this seems unlikely based on the successful isolation of other arenaviruses under similar conditions (Gómez et al., 2011). More likely is the possibility that samples contained viral titres that were too low for successful isolation in the cell lines tested (Lednicky et al., 2012) or that cell lines were contaminated with Mycoplasma. Isolation attempts were made in cell lines previously used to culture other mammarenaviruses (e.g. Vero, MDCK, BHK), but as Wēnzhōu virus was isolated in canine macrophage DH82 cell line (Li et al., 2014), it would be prudent to attempt isolation in this cell line in the future. A serological test highly specific to Wēnzhōu virus is also needed. As such, the specificity of the ELISA test developed during this study needs to be assessed against other mammarenaviruses and potentially optimised further, or an alternative strategy employed. In order to strengthen the case for Wēnzhōu virus as an agent of human disease, future surveys would ideally include a comparison of the seroprevalence in exposed and non-exposed populations, using such a Wēnzhōu virus-specific test. Although attempts were made during this study to obtain sera from individuals not native to the Southeastern Asian region, only limited samples were available. Many of these were from individuals who had been resident in this region for some time and therefore may have been exposed to Southeastern Asian mammarenaviruses. Sourcing and testing of sera from non-endemic areas is therefore needed. Studies aimed at detecting seroconversion in conjunction with molecular testing for viral RNA are also required, with the ultimate goal of isolating the virus in tissue culture or rodents. If Wēnzhōu virus does cause disease, it is also possible that like LASV and LCMV, more severe disease may occur in particular individuals such as pregnant women and immuno-compromised individuals (Bonthius, 2012; Price et al., 1988) and this too should be examined.

Globally infectious diseases remain a major cause of mortality and morbidity resulting in more than 15 million deaths per annum (Morens et al., 2004). However it is clear that many of the pathogenic agents responsible still remain undiagnosed, even when they are associated with potentially life threatening conditions. To improve protection we need a better understanding of both the ecology and epidemiology of diseases if we are to succeed in preventing them. Our findings illustrate how a 'One Health' approach in a regional hot-spot for emerging infectious disease can lead to the detection and characterisation of viruses likely associated with previously unrecognized human infections.

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Author details

1. Kim R Blasdell

   1. Institut Pasteur in Cambodia, Phnom Penh, Cambodia
   2. Commonwealth Scientific and Industrial Research Organisation, Australian Animal Health Laboratory, Geelong, Australia

   Contribution

   KRB, Conception and design, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article

   Contributed equally with

   Veasna Duong

   Competing interests

   No competing interests declared.

2. Veasna Duong

   Institut Pasteur in Cambodia, Phnom Penh, Cambodia

   Contribution

   VDu, Conception and design, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article

   Contributed equally with

   Kim R Blasdell

   Competing interests

   No competing interests declared.

3. Marc Eloit

   Institut Pasteur, Paris, France

   Contribution

   ME, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

   Competing interests

   No competing interests declared.

4. Fabrice Chretien

   Institut Pasteur, Paris, France

   Contribution

   FC, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

   Competing interests

   No competing interests declared.

5. Sowath Ly

   Institut Pasteur in Cambodia, Phnom Penh, Cambodia

   Contribution

   SL, Acquisition of data, Analysis and interpretation of data

   Competing interests

   No competing interests declared.

6. Vibol Hul

   Institut Pasteur in Cambodia, Phnom Penh, Cambodia

   Contribution

   VH, Conception and design, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

   Competing interests

   No competing interests declared.

7. Vincent Deubel

   Institut Pasteur in Cambodia, Phnom Penh, Cambodia

   Contribution

   VDe, Analysis and interpretation of data, Drafting or revising the article

   Competing interests

   No competing interests declared.

8. Serge Morand

   Institut des Sciences de l'Evolution, CNRS, IRD, Université Montpellier, Montpellier, France

   Contribution

   SM, Conception and design, Drafting or revising the article, Contributed unpublished essential data or reagents

   Competing interests

   No competing interests declared.

   "This ORCID iD identifies the author of this article:" 0000-0003-3986-7659

9. Philippe Buchy

   1. Institut Pasteur in Cambodia, Phnom Penh, Cambodia
   2. GlaxoSmithKline Vaccines R&D, Singapore, Singapore

   Contribution

   PB, Conception and design, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

   For correspondence

   buchyphilippe@hotmail.com

   Competing interests

   PB: Currently an employee of GSK Vaccines R&D in Asia-Pacific region

   "This ORCID iD identifies the author of this article:" 0000-0003-1372-3008

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