West Nile virus neuroinvasive disease incidence in the United States, 2002-2006

As the geographic range of reported human West Nile virus (WNV) disease has expanded across the United States, seasonal transmission and outbreaks have persisted over several years in many areas of the country. West Nile virus neuroinvasive disease (WNND) case reports from 2002 to 2006 were reviewed to determine which areas of the country have the highest reported cumulative incidence and whether those areas have had consistently high annual incidence. During the 5-year period examined, 9632 cases of WNND were reported nationwide. The cumulative incidence of WNND ranged from 0.2 to 32.2 per 100,000 population by state and from 0.1 to 241.2 per 100,000 population by county. States and counties with the highest cumulative incidence were primarily located in the northern Great Plains. States with consistently high annual incidence included South Dakota, North Dakota, Wyoming, New Mexico, Mississippi, Nebraska, Louisiana, and Colorado. All of these states, with the exception of New Mexico, were also among the states with the highest cumulative incidence. Counties with repeatedly high annual incidence were also primarily in the Great Plains and mid-South. The risk of WNND appears to be highest in areas where the primary WNV vectors are Culex tarsalis and Cx. quinquefasciatus mosquitoes.     

West Nile virus epizootiology in the southeastern United States, 2001

We investigated mosquito and bird involvement in West Nile virus (WNV) transmission in July 2001 in Jefferson County, FL, and Lowndes County, GA. We detected 16 WNV-infected pools from Culex quinquefasciatus, Cx. salinarius, Cx. nigripalpus, and Culiseta melanura. In Florida, 11% of 353 bird sera neutralized WNV. Antibody prevalence was greatest in northern cardinal (Cardinalis cardinalis, 75%), northern mockingbird (Mimus polyglottus, 50%), common ground-dove (Columbina passerina, 25%), common grackle (Quiscalus quiscula, 15%), domestic chicken (Gallus gallus, 16%), and house sparrow (Passer domesticus, 11%). Antibody-positive birds were detected in nine of 11 locations, among which prevalence in chickens ranged from 0% to 100%. Seropositive chickens were detected in Georgia as well. The primary transmission cycle of WNV in the southeastern United States apparently involves Culex mosquitoes and passerine birds. Chickens are frequently infected and may serve as effective sentinels in this region.     

Spread of equine West Nile virus encephalomyelitis during the 2002 Texas epidemic

Using reports of clinical West Nile virus (WNV) encephalomyelitis in Texas equids during 2002, the distribution of disease was analyzed using cluster statistics and spatial modeling to develop hypotheses of disease spread during the first year of its detection. Significant (P < 0.05) clusters of cases reported early during the outbreak were identified in east, northcentral, and north Texas, and significant (P < 0.05) clusters late during the outbreak were detected in central, south, and west Texas. Two counties on the south Texas coast first reported disease significantly (P < 0.05) earlier than their 10 nearest neighboring counties. The estimated incidence of disease was greatest in the high plains of north Texas and in northcentral Texas. Higher rates were also estimated in eastern and southern areas of the Gulf Coast. The spatial and temporal distribution observed indicates that the equine WNV epidemic began in two parts of Texas and spread elsewhere throughout the state. The mechanism of introduction and spread remains speculative.     

Inter-annual associations between precipitation and human incidence of West Nile virus in the United States

Higher-than-average precipitation levels may cause mosquito outbreaks if mosquitoes are limited by larval habitat availability. Alternatively, recent ecological research suggests that drought events can lead to mosquito outbreaks the following year due to changes in food web structure. By either mechanism, these mosquito outbreaks may contribute to human cases of West Nile Virus (WNV) in the recent United States outbreak. Using countylevel precipitation and human WNV incidence data (2002-2004), we tested the impacts of above and below-average rainfall on the prevalence of WNV in human populations both within and between years. We found evidence that human WNV incidence is most strongly associated with annual precipitation from the preceding year. Human outbreaks of WNV are preceded by above-average rainfall in the eastern United States and below-average rainfall in the western United States in the prior year. While no direct mechanism may be determined from this study, we hypothesize that differences in the ecology of mosquito vectors may be responsible for the opposite relationships between precipitation and WNV outbreaks between the eastern and western United States.     

Seroprevalence of West Nile virus in feral horses on Sheldon National Wildlife Refuge, Nevada, United States

We screened 1,397 feral horses (Equus caballus) on Sheldon National Wildlife Refuge, Nevada, United States, for IgM and IgG against flavivirus during 2004-2006, 2008, and 2009. Positive serum samples were tested for neutralizing antibodies to West Nile virus (WNV) and St. Louis encephalitis virus (SLEV). One animal was positive for antibody against WNV in 2004, but all others tested in 2004-2006 were negative. In 2008 and 2009, we found evidence of increasing seropositive horses with age, whereas seroprevalence of WNV decreased from 19% in 2008 to 7.2% in 2009. No horses were positive for antibody against SLEV. Being unvaccinated, feral horses can be useful for WNV surveillance.     

Culex flavivirus and West Nile virus mosquito coinfection and positive ecological association in Chicago, United States

Culex flavivirus (CxFV) is an insect-specific flavivirus globally distributed in mosquitoes of the genus Culex. CxFV was positively associated with West Nile virus (WNV) infection in a case-control study of 268 mosquito pools from an endemic focus of WNV transmission in Chicago, United States. Specifically, WNV-positive Culex mosquito pools were four times more likely also to be infected with CxFV than were spatiotemporally matched WNV-negative pools. In addition, mosquito pools from residential sites characterized by dense housing and impermeable surfaces were more likely to be infected with CxFV than were pools from nearby urban green spaces. Further, 6/15 (40%) WNV-positive individual mosquitoes were also CxFV positive, demonstrating that both viruses can coinfect mosquitoes in nature. Phylogenetic analysis of CxFV from Chicago demonstrated a pattern similar to WNV, consisting of low global viral diversity and lack of geographic clustering. These results illustrate a positive ecological association between CxFV and WNV, and that coinfection of individual mosquitoes can occur naturally in areas of high flaviviral transmission. These conclusions represent a challenge to the hypothesis of super-infection exclusion in the CxFV/WNV system, whereby an established infection with one virus may interfere with secondary viral infection with a similar virus. This study suggests that infection with insect-specific flaviviruses such as CxFV may not exclude secondary infection with genetically distinct flaviviruses such as WNV, and that both viruses can naturally coinfect mosquitoes that are epidemic bridge vectors of WNV to humans.     

Early season crow mortality as a sentinel for West Nile virus disease in humans,northeastern United States

The 1999 New York epidemic of human West Nile virus (WN) encephalitis and meningitis was preceded by a crow die-off also caused by WN infection. As one component of the subsequently developed national surveillance system, crow mortality data were collected to detect WN activity before humans might become infected. However, predicting areas at risk for human WN disease likely requires assessment of multiple factors, including the intensity and timing of crow epizootics. To identify early season measures of WN activity in crows associated with subsequent WN disease in humans, county-level crow mortality data from seven northeastern states were analyzed. A predictive model was developed based on analysis of 2000 surveillance data and then assessed for 2001. To characterize the intensity of early season WN activity in crows, 15 variables were constructed from surveillance data of 52 counties that tested at least four crows during the early season (defined as June 17-July 28, 2000). County values for each variable were dichotomized at the 75th percentile into "high" and "low" activity. Multivariate analysis indicated that "high" early season activity of two variables-density of reported dead crow sightings (reported dead crows/area) and [(WN-infected crows/tested crows) x (human population)]--were associated with report of at least one human WN disease case (for each variable: adjusted odds ratio, 6.9; 95% confidence interval, 1.2-40.6). An assessment of this model using 2001 surveillance data from 61 counties yielded similar findings. With emphasis on early season WN activity, crow surveillance may allow timely targeting of interventions to protect the public health.     

West Nile encephalitis: an emerging disease in the United States.

In 1999, an epidemic of West Nile virus (WNV) encephalitis occurred in New York City (NYC) and 2 surrounding New York counties. Simultaneously, an epizootic among American crows and other bird species occurred in 4 states. Indigenous transmission of WNV had never been documented in the western hemisphere until this epidemic. In 2000, the epizootic expanded to 12 states and the District of Columbia, and the epidemic continued in NYC, 5 New Jersey counties, and 1 Connecticut county. In addition to these outbreaks, several large epidemics of WNV have occurred in other regions of the world where this disease was absent or rare >5 years ago. Many of the WNV strains isolated during recent outbreaks demonstrate an extremely high degree of homology that strongly suggests widespread circulation of potentially epidemic strains of WNV. The high rates of severe neurologic illness and death among humans, horses, and birds in these outbreaks are unprecedented and unexplained. We review the current status of WNV in the United States.     

Importance of vertical and horizontal transmission of West Nile virus by Culex pipiens in the Northeastern United States

West Nile virus (WNV) has become established in the northeastern United States, where mosquitoes are inactive during winter. There have been no documented studies to explain how this virus survives winter and reinitiates infection in spring. We report that WNV was vertically transmitted to 2 F(1) female Culex pipiens from a naturally infected female collected in Stratford, Connecticut. One vertically infected F(1) female, which was 168 days old, fed on a hamster that died 8 days later of West Nile disease. This suggests that WNV survives winter in unfed, vertically infected C. pipiens with amplification initiated in spring by horizontal transmission.     

Host feeding patterns of established and potential mosquito vectors of West Nile virus in the eastern United States

An important variable in determining the vectorial capacity of mosquito species for arthropod-borne infections is the degree of contact of the vector and the vertebrate reservoir. This parameter can be estimated by examining the host-feeding habits of vectors. Serological and polymerase chain reaction based methods have been used to study the host-feedings patterns of 21 mosquito species from New York, New Jersey, and Tennessee, 19 of which previously have been found infected with West Nile virus. Mammalophilic mosquito species in New Jersey and New York fed primarily upon white-tailed deer, while those from Memphis, Tennessee, fed mainly upon domestic dogs. A total of 24 different avian host species were detected among the avian-derived blood meals. American Robin, Northern Cardinal, Northern Mockingbird, Tufted Titmouse, and Brown-headed Cowbird were common avian hosts, while blood meals derived from the American Crow were relatively rare. Although the majority of common host species were potentially among the most abundant birds at each location, the proportion of blood meals from the most commonly fed upon avian species was greater than was predicted based upon the likely abundance of these species alone. These findings suggest that vector species for West Nile virus may preferentially feed upon certain avian hosts.     

Avian hosts for West Nile virus in St. Tammany Parish, Louisiana, 2002

West Nile virus (WNV) infections in free-ranging birds were studied in Slidell, St. Tammany Parish, Louisiana, after a human encephalitis outbreak peaked there in July 2002. Seroprevalence in resident, free-ranging wild birds in one suburban site was 25% and 24% in August and October, respectively, indicating that most transmission had ceased by early August. Mortality rates, seroprevalence rates, host competence, and crude population estimates were used in mathematical models to predict actual infection rates, population impacts, and importance as amplifying hosts for several common passerine birds. Northern cardinal (Cardinalis cardinalis) and house sparrow (Passer domesticus) were the principal amplifying hosts, but blue jay (Cyanocitta cristata) and northern mockingbird (Mimus polyglottos) also contributed. The blue jay population was reduced by an estimated 47%. A variety of passerine bird species combined to play an important role as amplifying hosts in the WNV transmission cycle.     

Differential West Nile fever ascertainment in the United States: a multilevel analysis

We evaluated the completeness of West Nile fever (WNF) surveillance within the U.S. public health system. We surveyed laboratory and surveillance programs on policies, practices, and capacities for testing, confirmation, and reporting (collectively called ascertainment) from 2003 through 2005. We calculated syndrome ascertainment ratios by dividing WNF counts by neuroinvasive disease counts; separately, we performed multilevel modeling. Jurisdictions were more likely to ascertain at least one WNF cases per West Nile neuroinvasive disease case when ≤ 1 testing restrictions existed (odds ratio [OR] = 7.7, 95% confidence interval [CI] = 1.3-46.4), when conducting ≥ 4 activities to enhance reporting (OR = 9.3, 95% CI = 1.6-54.8), and when ≥ 5.0 staff per million residents were dedicated to arboviral surveillance (OR = 6.4, 95% CI = 1.0-40.3). Ascertainment of WNF was less likely among Blacks (OR = 0.56, 95% CI = 0.31-0.99) and Hispanics (OR = 0.69, 95% CI = 0.48-0.98) than among Whites. Ascertainment was more complete when testing and reporting were enhanced, but differentially incomplete for minorities.     

West Nile virus

West Nile (WN) virus is a mosquito-borne flavivirus and human, equine, and avian neuropathogen. The virus is indigenous to Africa, Asia, Europe, and Australia, and has recently caused large epidemics in Romania, Russia, and Israel. Birds are the natural reservoir (amplifying) hosts, and WN virus is maintained in nature in a mosquito-bird-mosquito transmission cycle primarily involving Culex sp mosquitoes. WN virus was recently introduced to North America, where it was first detected in 1999 during an epidemic of meningoencephalitis in New York City. During 1999-2002, the virus extended its range throughout much of the eastern parts of the USA, and its range within the western hemisphere is expected to continue to expand. During 1999-2001, 142 cases of neuroinvasive WN viral disease of the central nervous system (including 18 fatalities), and seven cases of uncomplicated WN fever were reported in the USA. Most human WN viral infections are subclinical but clinical infections can range in severity from uncomplicated WN fever to fatal meningoencephalitis; the incidence of severe neuroinvasive disease and death increase with age. Serology remains the mainstay of laboratory diagnosis. No WN virus-specific treatment or vaccine is available. Prevention depends on organised, sustained vector mosquito control, and public education.     

Regional differences in the association between land cover and West Nile virus disease incidence in humans in the United States

West Nile virus (WNV) is generally considered to be an urban pathogen in the United States, but studies associating land cover and disease incidence, seroprevalence, or infection rate in humans, birds, domesticated and wild mammals, and mosquitoes report varying and sometimes contradictory results at an array of spatial extents. Human infection can provide insight about basic transmission activity; therefore, we analyzed data on the incidence of WNV disease in humans to obtain a comprehensive picture of how human disease and land cover type are associated across the United States. Human WNV disease incidence in Northeastern regions was positively associated with urban land covers, whereas incidence in the Western United States was positively associated with agricultural land covers. We suggest that these regional associations are explained by the geographic distributions of prominent WNV vectors: Culex pipiens complex (including Cx. pipiens and Cx. quinquefasciatus) in the Northeast and Cx. tarsalis in the Western United States.     

Regional Variation of Climatic Influences on West Nile Virus Outbreaks in the United States

The national resurgence of human West Nile virus (WNV) disease in 2012 raised questions about the factors responsible for WNV outbreaks. Interannual climatic variations may influence WNV amplification and transmission to humans through multiple pathways, including mosquito breeding habitats, gonotrophic cycles, extrinsic incubation, avian communities, and human behavior. We examined the influences of temperature and precipitation anomalies on interannual variation in human WNV cases in three regions of the United States. There were consistent positive influences of winter temperatures, weaker and more variable positive effects of spring and summer temperatures, and highly variable precipitation effects that ranged from positive to negative. The overwintering period may be a particularly important climatic constraint on the dynamics of WNV in cold-temperate regions of North America. Geographic differences in the seasonal timing and relative importance of climatic drivers of WNV risk likely reflect underlying variability in key ecological and social characteristics.     

Remotely-sensed vegetation indices identify mosquito clusters of West Nile virus vectors in an urban landscape in the northeastern United States

Heterogeneity in urban landscapes can influence the effectiveness of mosquito-borne disease control. We used remotely sensed vegetation indices to discriminate among mosquito habitats within a densely populated urban environment in New Haven, CT. ASTER derived vegetation indices were identified for 16 sites where adult mosquitoes were trapped over the summer of 2004. Canonical correlation analysis showed a significant relationship between the environmental variables (normalized difference vegetation index, disease/water stress index and distance to water) and four local West Nile virus competent vectors (Cx. pipiens, Cx. restuans, Cx. salinarius, and Ae. vexans) (0.93, P = 0.03) explaining 86% of the variance in the environmental and mosquito measures. Sites were clustered based on these remotely sensed environmental variables. Three clusters were identified which provide insight into the distribution of West Nile virus vectors in an urban area. Identification of habitat differences of mosquitoes within the urban landscape has important implications for understanding West Nile virus transmission and for control of vector-competent mosquito species.     

The traveller and West Nile virus

Infection with West Nile virus is a zoonosis which has emerged recently as a threat to public health and animal health in temperate regions in parts of Europe and more extensively in North America. Most infections are asymptomatic, 20% lead to a mild febrile illness and about 1% result in severe neurological disease, particularly in those over the age of 50 years. West Nile virus is transmitted by mosquitoes and the principal hosts are numerous species of wild birds. There is no specific treatment and a vaccine is not available. Prevention in areas where the virus is circulating is based on protection from mosquito bites and reduction and control of the number of mosquitoes in the environment. The risk to travellers is not known at present, but appears to be low in terms of symptomatic illness.