Emergence and persistence of hantaviruses.

Hantaviral diseases have been recognized for hundreds of years but, until 1976, they had not been associated with an infectious agent. When Lee and colleagues isolated what is now known as Hantaan virus, the techniques they introduced allowed further investigations into the etiology of the classical hantavirus disease, hemorrhagic fever with renal syndrome (HFRS), now known to be caused by any of multiple hantaviruses. The discovery of hantavirus pulmonary syndrome (HPS) in the New World, and that it also can be caused by any of multiple hantaviruses (family Bunyaviridae, genus Hantavirus), has opened an entire field of epidemiologic, virologic, molecular, behavioral, and ecologic studies of these viruses. There appears to be a single hantavirus-single rodent host association, such that understanding the idiosyncrasies of each rodent host species and the ecologic variables that affect them are recognized as critical if we are to reduce human risk for infection. This chapter summarizes what is known about hantaviruses with regard to history of these viruses, their taxonomy, recognized geographical distribution, ecologic factors impacting their maintenance and spread of hantaviruses, effect of rodent behavior on hantavirus transmission, influence of host factors on susceptibility to and transmission of hantaviruses, and transmission of hantaviruses from rodents to humans. In addition, we summarize all these complexities and provide suggestions for future research directions.

Transmission of an arenavirus in white-throated woodrats (Neotoma albigula), southeastern Colorado, 1995-1999.

From 1995 to 1999, we conducted longitudinal studies of white- throated woodrats (Neotoma albigula) in southeastern Colorado. Forty-five (42.9%) of 105 female and 15 (26.8%) of 56 male N. albigula had antibodies against Whitewater Arroyo virus (WWAV). Sixteen female and three male N. albigula seroconverted during the study period, most of them during July-November, when population densities are highest. Analyses of longevity data, minimum numbers alive and infected, movements, and weight data suggest that the dominant mode of WWAV transmission among white-throated woodrats in Colorado is direct contact. WWAV was recently reported to cause fatal infection in humans. Our findings will lead to better assessment of the public health threat posed by infected woodrats and may be useful in predicting periods of increased risk for human infection.

Do unusual site-specific population dynamics of rodent reservoirs provide clues to the natural history of hantaviruses?

Between January 1995 and November 1997, longitudinal mark-recapture studies of rodent hosts of hantaviruses in a disturbed microhabitat within a shortgrass prairie ecosystem in southeastern Colorado (USA) were conducted. The site was distinguished by edaphic and floristic characteristics unique to this area and associated with historical land use patterns, as well as the year-around availability of water from a functioning windmill. Populations of two common rodent species that are hosts for hantaviruses, Peromyscus maniculatus and Reithrodontomys megalotis, had unusually rapid turnover, a younger age structure, and a much lower prevalence of antibody to Sin Nombre virus than did populations at nearby sites in more typical shortgrass prairie and canyon habitats. Based on these findings, we suggest that a stable resident population of the reservoir is critical to the maintenance of hantaviruses at a given site, and we hypothesize that long-lived, persistently infected rodents are the principal transseasonal reservoir of hantaviruses.

Dual captures of Colorado rodents: implications for transmission of hantaviruses.

We analyzed dual-capture data collected during longitudinal studies monitoring transmission and persistence of Sin Nombre virus in rodents in Colorado. Our data indicate that multiple captures (two or more rodents captured in a single trap) may not be random, as indicated by previous studies, but rather the result of underlying, species-specific social behavior or cohesiveness. In the pairs we captured, most often, rodents were of the same species, were male, and could be recaptured as pairs. Therefore, dual captures of rodents, which are unusual but not rare, tend to occur among certain species, and appear to be nonrandom, group-foraging encounters. These demographic and ecologic characteristics may have implications for the study of the transmission of hantaviruses.

Relationships of deer mouse movement, vegetative structure, and prevalence of infection with Sin Nombre virus.

The effects of vegetative structure on movement of deer mice (Peromyscus maniculatus) were examined in two distinct vegetation associations, one near Hesperus and the other near Molina in western Colorado (USA) from June-October 1994 to October 1998. We monitored movement by live-trapping small mammals within Gambel's oak/mixed-grass (Hesperus) and sage brush/juniper (Molina) vegetation types. Vegetative structure differed between the sites with Molina having more cover provided by shrubs and Hesperus having more cover provided by forbs. Adult male deer mice moved greater distances at Hesperus than at Molina. Sub-adult males tended to move greater distances than did adult females. Relative abundances of deer mice tended to differ by season, but the average relative abundance of deer mice was greater at Molina. Long-term prevalence of infection with SNV was greater at Hesperus and was greatest in adult males at Hesperus (36.1%). Adult males at Molina exhibited a prevalence of infection with SNV of 25.0%. Infection with SNV was highly associated with scars or wounds for adult male, adult female, and juvenile male deer mice at Hesperus, but only for adult female deer mice at Molina. The presence of scars or wounds tended to be associated with greater age, but male deer mice at Hesperus were more likely to have wounds than female deer mice of the same age class. A similar pattern, excluding juveniles, was observed at Molina. Intraspecific interactions and environmentally elicited long-distance movements of deer mice may play a role in prevalence of infection with SNV in these animals.

Natural history of Sin Nombre virus in western Colorado.

A mark-recapture longitudinal study of immunoglobulin G (IgG) antibody to Sin Nombre virus (SNV) in rodent populations in western Colorado (1994-results summarized to October 1997) indicates the presence of SNV or a closely related hantavirus at two sites. Most rodents (principally deer mice, Peromyscus maniculatus, and pinyon mice, P. truei) did not persist on the trapping webs much beyond 1 month after first capture. Some persisted more than 1 year, which suggests that even a few infected deer mice could serve as transseasonal reservoirs and mechanisms for over-winter virus maintenance. A positive association between wounds and SNV antibody in adult animals at both sites suggests that when infected rodents in certain populations fight with uninfected rodents, virus amplification occurs. At both sites, male rodents comprised a larger percentage of seropositive mice than recaptured mice, which suggests that male mice contribute more to the SNV epizootic cycle than female mice. In deer mice, IgG antibody prevalence fluctuations were positively associated with population fluctuations. The rates of seroconversion, which in deer mice at both sites occurred mostly during late summer and midwinter, were higher than the seroprevalence, which suggests that the longer deer mice live, the greater the probability they will become infected with SNV.

Taxonomy of African horse sickness viruses.

Nine distinct genera are currently recognised within the virus family Reoviridae, which include a total of 63 virus groups or species (species = virus group = electropherotype or serogroup), comprising 214 virus serotypes or subtypes, as well as 20 provisional types or subtypes, most of which (149 + 9 tentative) are assigned to the genus Orbivirus [5, 9, 16]. The 19 species of orbiviruses (serogroups), were established principally on antigenic (serologic) grounds but many of these placements have been supported by molecular analyses. This introductory paper defines the taxonomy and classification of these viruses and establishes guidelines for use in other paper to be presented at this symposium and elsewhere.

[Twenty-eight alphavirus strains isolated from mosquitoes and ticks captured from Hainan Island, China].

Virus isolation was carried out from mosquitoes and ticks captured in nature from Hainan Island of China from 1983-1988. Twenty-eight non-dengue and non-Japanese encephalitis viral strains were isolated from different places, of which 27 strains were recovered from different species of mosquitoes and 1 strain from ticks. At the Division of Vector-Bore Viral Diseases, National Center for Infectious Diseases, Centers for Disease Control, USA, identification of virus family and virus genus on these 28 viral strains was conducted by the use of a complete set of internationally standardized antibodies against arboviruses and by means of indirect immunofluorescence. Results proved that all these 28 viral strains belonged to the Genus Alphavirus of the Family Togaviridae.

Telford H. Work--a tribute.

The major contributions of Telford H. Work to the field of arbovirology and epidemiology of arthropod-transmitted diseases are detailed. The human aspects of his life are mentioned in this memorial address.

Typing of LaCrosse, snowshoe hare, and Tahyna viruses by analyses of single-strand conformation polymorphisms of the small RNA segments.

Single-strand conformation polymorphism analysis was developed to differentiate the small RNA segments of three California serogroup bunyaviruses. The small RNA segments of La Crosse, snowshoe hare, and Tahyna viruses were reverse transcribed and PCR amplified. The cDNAs were then denatured, rapidly chilled to promote intrastrand reassociation, separated electrophoretically on a nondenaturing gel at room temperature, and silver stained. The resulting single-strand conformation polymorphism patterns were specific for the respective viruses. This molecular technique offers great potential for virus typing and taxonomic studies.

Experimental infection of wading birds with eastern equine encephalitis virus.

To study the susceptibility of wading birds to eastern equine encephalitis (EEE) virus and to determine their potential as reservoir or amplifying hosts, fledgling glossy ibises (Plegadis falcinellus) and snowy egrets (Egretta thula) were captured in New Jersey (USA) and shipped to Colorado (USA) where they were experimentally inoculated with EEE virus. All 16 snowy egrets and 14 (93%) of 15 of the glossy ibises inoculated became viremic with moderate titers, and all survivors developed neutralizing antibody. Six ibises and two egrets died during the first week after inoculation, and EEE virus was isolated from the tissues of three birds. Our experimental results support field evidence about the relative involvement of glossy ibises and snowy egrets in the epizootiology of EEE virus in New Jersey.