Low oral receptivity for dengue type 2 viruses of Aedes albopictus from Southeast Asia compared with that of Aedes aegypti

Dengue hemorrhagic fever has been a major health problem in Asia since the 1950s. During this period, the former principal vector of dengue viruses in Asia, Aedes albopictus, was replaced by Aedes aegypti in most major cities of the area. Ae. aegypti is now considered the main vector of dengue viruses in Asia. Surprisingly, however, this mosquito has been described as having a relatively low oral receptivity for dengue viruses compared with Ae. albopictus. In the present study, we compared the relative oral receptivities of Ae. aegypti and Ae. albopictus collected in southeast Asia from both sympatric and allopatric breeding sites. In all instances, the oral receptivity of Ae. aegypti to the dengue type 2 virus used was significantly higher than that of Ae. albopictus. We also compared the relative oral receptivity of Ae. aegypti and Ae. albopictus for two other low-passage strains of dengue 2. In all instances, Ae. aegypti was significantly more receptive than Ae. albopictus. It should be noted, however, that the difference was found only for Ae. albopictus recently collected from the field (Ta Promh strain, Cambodia, 2001) and not for an Ae. albopictus strain that had been colonized for many years (Oahu strain, Hawaii, 1971). We also observed a significant increase in the infection rate of Ae. albopictus of the Ta Promh strain with increasing generations in the laboratory. These observations demonstrate the importance of considering the colonization history of mosquitoes when assessing their susceptibility to infection with dengue viruses and, perhaps, other arboviruses.     

Transovarial transmission of dengue viruses by mosquitoes: Aedes albopictus and Aedes aegypti

Transovarial transmission of all four dengue serotypes was demonstrated in Aedes albopictus mosquitoes. The rates of such transmission varied with the serotype and strain of virus. In general, the highest rates were observed with strains of dengue type 1 and the lowest with dengue type 3. Surprisingly, despite the use of viral strains of the four dengue serotypes which gave the highest rates with Ae. albopictus, transovarial transmission was observed in Aedes aegypti only with dengue type 1, and then only at a relatively low rate. Five different strains of Ae. aegypti were employed, including one that was known to be relatively susceptible to oral infection with dengue viruses. The findings support the view that Ae. aegypti, while of major importance from the point of view of transmission of dengue to man, may be relatively unimportant in the overall natural history of dengue viruses.     

A mosquito densovirus infecting Aedes aegypti and Aedes albopictus from Thailand.

A previously undescribed mosquito densovirus was detected in colonies of Aedes aegypti and Ae. albopictus from Thailand, using a polymerase chain reaction (PCR)-based assay. Phylogenetic analysis of this virus showed it to be most closely related to ADNV isolated from Russian Ae. aegypti. Both Aedes species were susceptible to oral infection with the Thai-strain virus. Larval mortality for Ae. albopictus was higher (82%) than for Ae. aegypti (51%). Aedes aegypti were able to transmit the virus vertically to a high (58%) proportion of G1 progeny, and the virus was maintained persistently for up to six generations. A PCR survey of adult Ae. aegypti and Ae. albopictus in Thailand indicated that only Ae. aegypti are infected in the field, with an overall prevalence of 44%. Densovirus infection in adult Ae. aegypti showed distinct seasonal variation. The Thai strain densovirus may play a role in structuring Ae. albopictus and Ae. aegypti populations in nature.     

Polygamy: the possibly significant behavior of Aedes aegypti and Aedes albopictus in relation to the efficient transmission of dengue virus

The polygamous behavior of male Aedes aegypti (L.) and Ae. albopictus (Skuse) was investigated by co-habiting a newly-emerged male and females in a 30 cm3 cage (1 male: 20 females) for up to 5 consecutive days. As determined by insemination rates, the results indicated that one Ae. aegypti and Ae. albopictus male could successfully mate with 1.10 (0-4), 4.10 (1-8), 5.40 (4-8), 5.10 (2-8), 5.15 (3-9) and 0.20 (0-3), 1.70 (0-3), 2.35 (1-4), 2.30 (0-4), 2.35 (1-4) Ae. aegypti and Ae. albopictus females, respectively on day 1,2,3,4 and 5 consecutively. The possibly significant role of their polygamy in relation to dengue virus transmission is discussed.     

A survey of dengue viral infection in Aedes aegypti and Aedes albopictus from re-epidemic areas in the north of Thailand using nucleic acid sequence based amplification assay

Immature stages of Aedes aegypti and Ae. albopictus were collected from 17 dengue re-epidemic areas in Chiang Mai and Lampang Provinces, in the north of Thailand. They were reared to adults and tested for dengue viral RNA by a nucleic acid sequence based amplification assay (NASBA). Of a total of 9,825 Ae. aegypti and 150 Ae. albopictus examined, none of them were found positive for the virus, suggesting that transovarial transmission may be very low in the vector populations and may not play a significant role in the epidemiology of dengue infection in Thailand.     

Dengue vector prevalence and virus infection in a rural area in south India

We conducted a 2-year (1997-1999) longitudinal, entomological and virological study in three dengue endemic villages in Vellore district, Tamil Nadu, to understand the dynamics of dengue transmission. Aedes aegypti (Linn.), Ae. albopictus (Skuse) and Ae. vittatus (Bigot) were the prevalent vector species. Aedes aegypti was breeding throughout the year with a Breteau index ranging from 9.05 to 45.49. Aedes albopictus and Ae. vittatus were prevalent mainly in the rainy season. Small water holding containers (cemented tanks/cisterns) were the perennial breeding source of Ae. aegypti, and its abundance was significantly higher in semi-urbanized central areas than the peripheral areas of the villages. From 271 pools (4016 specimens) of adult females, eight dengue virus (DENV) isolates were obtained of which seven were from Ae. aegypti and one from Ae. albopictus. This is the first report of DENV isolation from Ae. albopictus in rural India. Infection rates in the two species were comparable. However, due to higher and perennial prevalence, Ae. aegypti is considered as primary vector with Ae. albopictus playing a secondary role. Despite circulation of all four serotypes (DENV 1-4) detected mainly during the transmission season, the high anthropophilic index of the vectors and their abundance, no human dengue case was reported, suggesting silent dengue transmission.     

Growth characteristics of ChimeriVax-DEN2 vaccine virus in Aedes aegypti and Aedes albopictus mosquitoes

The chimeric yellow fever (YF) 17D-dengue type 2 (ChimeriVax-DEN2) vaccine virus developed by Acambis, Inc. (Cambridge, MA) contains the prM and E genes of wild-type (wt) dengue 2 (DEN-2) (strain PUO-218) virus in the YF vaccine virus (strain 17D) backbone. The potential of ChimeriVax-DEN2 virus to infect and be transmitted by Aedes aegypti, the principal DEN and YF virus mosquito vector, and Aedes albopictus, a species that occurs in areas of active transmission of YF and DEN viruses, was evaluated. Mosquitoes were intrathoracically (IT) inoculated with virus or were fed a virus-laden blood meal, and the replication kinetics of ChimeriVax-DEN2 were compared with the wt DEN-2 and YF 17D vaccine viruses. Replication of YF 17D virus is attenuated in cultured Ae. albopictus C6/36 mosquito cells and in Ae. aegypti and Ae. albopictus mosquitoes. Growth of ChimeriVax-DEN2 virus similarly was restricted in C6/36 cells and in mosquitoes. ChimeriVax-DEN2 replicated in 56% of IT inoculated Ae. aegypti, and virus disseminated to head tissue in 36%, with a mean viral titer of 1.8 log10 PFU/mosquito. Of mosquitoes, 16% of Ae. aegypti and 24% of Ae. albopictus were infected 14 days after a blood meal containing ChimeriVax-DEN2, but virus did not disseminate to head tissue. In contrast, DEN-2 replicated in all IT inoculated and orally infected Ae. aegypti (mean titer 5.5 log10 PFU/mosquito), and virus disseminated to head tissue in 95%. Of Ae. albopictus, 84% were infected after a blood meal containing DEN-2 virus; dissemination occurred in 36%. Replication of ChimeriVax-DEN2 virus in mosquitoes corresponded to that of YF 17D vaccine virus, which is restricted in its ability to infect and replicate in mosquitoes. Therefore, transmission of ChimeriVax-DEN2 virus by vector mosquitoes is unlikely.     

Infection of adult Aedes aegypti and Ae. albopictus mosquitoes with the entomopathogenic fungus Metarhizium anisopliae

This study describes a laboratory investigation on the use of the insect-pathogenic fungus Metarhizium anisopliae against adult Aedes aegypti and Ae. albopictus mosquitoes. At a dosage of 1.6 x 10(10)conidia/m(2), applied on material that served as a mosquito resting site, an average of 87.1+/-2.65% of Ae. aegypti and 89.3+/-2.2% of Ae. albopictus became infected with the fungus. The life span of fungus-contaminated mosquitoes of both species was significantly reduced compared to uninfected mosquitoes. LT(50)-values of fungus-contaminated mosquitoes ranged between 3.1+/-0.2 days (male Ae. aegypti) and 4.1+/-0.3 days (female Ae. aegypti). LT(50)-values of uncontaminated mosquitoes ranged from 17.7+/-0.4 days (female Ae. albopictus) to 19.7+/-0.6 days (male Ae. albopictus). These results indicate that both mosquito species are highly susceptible to infection with this entomopathogen. Requirements for developing and incorporating this biological control method into current strategies to control major diseases vectored by these species, such as dengue fever, are discussed.     

Distribution and seasonality of vertically transmitted dengue viruses in Aedes mosquitoes in arid and semi-arid areas of Rajasthan, India

BACKGROUND & OBJECTIVES: Transovarial transmission of dengue virus is a crucial etiological phenomenon responsible for persistence of virus during inter-epidemic period of the disease. Distribution and seasonality of this phenomenon in disease endemic areas may contribute to explain emergence of dengue and its subsequent prevention. The study on seasonal and area distribution of transovarial transmission of dengue virus in Aedes aegypti, Ae. albopictus and Ae. vittatus has been made in desert and non-desert districts of Rajasthan, India from 2006 to 2007. The observations revealed role of different Aedes species in transmission and retention of dengue virus. METHODS: The larvae of Ae. aegypti, Ae. albopictus and Ae. vittatus were collected during each of the study seasons from rural and urban areas of three districts-Jodhpur, Jaipur and Kota. The larvae were collected from domestic and peri-domestic containers and from tree holes of peri-urban foci such as gardens and parks and were reared into adults in the laboratory at room temperature. The laboratory reared adults were subjected to Indirect fluorescence antibody test (IFAT). The laboratory-reared adult mosquitoes showing positive IFA were treated as the sample showing vertically transmitted dengue virus. RESULTS: Pooled data for all the four seasons revealed maximum (15.7%) mosquito infectivity in Ae. albopictus followed by Ae. aegypti (12.6%) in Jodhpur district. In Jaipur district, Ae. vittatus showed highest infection (20%) of vertically transmitted virus followed by Ae. albopictus (18.7%) and least in Ae. aegypti (13.3%). In Kota district, pooled data for all the four seasons showed maximum vertical infection of mosquitoes in Ae. albopictus (14.2%). INTERPRETATION & CONCLUSION: Transovarial transmission of dengue virus by available vector species in a dengue endemic setting could be the key etiological phenomenon responsible for re-emergence of the disease from inter-epidemic to epidemic phase of disease onset. The observations in the present study suggest that during winter season which is not the active transmission season of dengue in Rajasthan, Ae. albopictus has shown maximum percentage of vertically transmitted virus. Our observation substantiates with the earlier studies that how Ae. albopictus is horbouring virus during inter-epidemic period of dengue. Another important lead emerging through present study is the high mosquito infectivity of Ae. aegypti during summer and rainy seasons especially from desert districts, Jodhpur and semi district. This observation suggests that in Rajasthan, owing to tendency of overstorage of domestic water by the inhabitants, mosquito and vertically transmitted virus get pronounced during summer season which could precedes the active transmission season of dengue during following rainy season.     

Use of a geographic information system for defining spatial risk for dengue transmission in Bangladesh: role for Aedes albopictus in an urban outbreak

We used conventional and spatial analytical tools to characterize patterns of transmission during a community-wide outbreak of dengue fever and dengue hemorrhagic fever in Dhaka, Bangladesh in 2000. A comprehensive household-level mosquito vector survey and interview was conducted to obtain data on mosquito species and breeding as well as illness consistent with dengue. Clusters of dengue illnesses and high-density vector populations were observed in a distinct sector of the city. Dengue clusters are less identifiable in areas further away from major hospitals, suggesting that proximity to hospitals determines whether cases of dengue are diagnosed. Focusing on those areas relatively close to hospitals, we found a spatial association between dengue clusters and vector populations. Households reporting a recent dengue illness were more likely to have Aedes albopictus larvae present in the home when compared with households not reporting cases. Households reporting a recent dengue illness were also more likely to have a neighbor with Ae. albopictus present in the home. In contrast, the presence of Aedes aegypti within the premises as well as the homes of neighbors (within 50 meters) was not associated with dengue illness. Given that the breeding habitats for Ae. albopictus are somewhat distinct from those of Ae. aegypti, the findings of this study have implications for control of dengue transmission in this urban setting where much of the focus has been on indoor mosquito breeding and transmission. Public health officials may find the disease-environment map useful for planning targeted interventions because it displays areas where transmission is most intense.     

Role of Aedes aegypti and Aedes albopictus during the 2011 dengue fever epidemics in Hanoi,Vietnam

Objective: To record the human cases of dengue fever(DF) and investigate the Aedes mosquito species circulating during the Hanoi 2011 DF epidemics. Methods: 24 different outbreak points were recorded in 8 districts between August and December 2011. Results: 140 patients were hospitalized following dengue diagnostic with a predominance of males(59.3%) and the 15-34 age class. Only DENV-1(11.27%) and DENV-2(88.73%) serotypes were detected in human samples. Mosquito sampling performed in and around patients households revealed the predominance of Aedes aegypti(95.15%) versus Aedes albopictus(4.85%). There is a positive correlation between the population density of Aedes aegypti and the number of human cases and duration of outbreaks. Conclusions: This was not observed for Aedes albopictus. 3 pools of Aedes aegypti were positive with dengue virus, two with DENV-1 and one with DENV-2.     

Seasonality and insecticide susceptibility of dengue vectors: an ovitrap based survey in a residential area of northern Sri Lanka

With the backdrop of a high incidence of dengue in Jaffna District, Sri Lanka, an ovitrap based survey was carried out from May 2005 to April 2006 in a residential area to study the seasonality and insecticide susceptibility of Aedes aegypti and Ae. albopictus. Conventional ovitraps were placed inside and outside of 10 randomly selected houses to collect indoor breeding and outdoor breeding Aedes mosquitoes; collections took place fortnightly. A total of 3075 Ae. aegypti and 2665 Ae. albopictus were collected in outdoor ovitraps, whereas in indoor ovitraps a total of 2528 Ae. aegypti and 2002 Ae. albopictus were collected. The highest values for Aedes density and positive ovitrap percentage were recorded in January 2006. A seasonal shift was observed in the density of Ae. aegypti and Ae. albopictus. Ae aegypti density was high during and after the Northeast monsoon whilst Ae. albopictus was the dominant species during the onset of the Northeast monsoon. A significant association (p < 0.05) between Aedes density and rainfall was observed. The association of these two species to site, either indoors or outdoors, was not statistically significant (p > 0.05). Both the species were found to be highly resistant to 4% DDT and completely susceptible to 5% malathion. The high prevalence and the ability of both species to breed indoors and outdoors should be taken into account when formulating a dengue vector control program with community participation in the Jaffna District, Sri Lanka.     

Epidemic dengue hemorrhagic fever in rural Indonesia. III. Entomological studies.

Entomological studies were carried out during a dengue hemorrhagic fever epidemic in Central Java in December 1976. Both Aedes aegypti and Aedes albopictus were widely distributed in Bantul, but the latter species was more abundant. Comparative studies on the vector competence of the two species showed that Ae. albopictus had a higher susceptibility than Ae. aegypti to oral infection with all four dengue serotypes. The two species were equally compentent in transmitting the Bantul strain of dengue 3 virus after parenteral infection. The data suggest that Ae. albopictus could have been an important vector in this epidemic, but no direct observations were obtained to define the respective contribution of either Ae. albopictus or Ae. aegypti.     

Larval habitats and distribution patterns of Aedes aegypti (Linnaeus) and Aedes albopictus (Skuse), in Thailand

This study was conducted to survey larval breeding habitats and to obtain larval abundance during the dry period covering all 5 geographical zones of Thailand. Our results indicated Aedes aegypri is prevalent all over the country, whereas Aedes albopictus is more restricted to the remote area of the south. Water storage containers, especially water jars, served as a main larval breeding habitats of Ae. aegypti, whereas broken cans and plastic containers are considered primary breeding sites for Ae. alpopictus during the dry period. In addition, Aedes larval indices, container index (CI), house index (HI), and Breteau index (BI) were measured. CI and HI values from the central part were significantly higher than those from other areas (p<0.01). BI values of all collection sites were greater than 50 (a maximum BI value accepted by the Ministry of Public Health, Thailand). In brief, Ae. aegypti and Ae. albopictus populations heavily infested many towns and residential areas of the country. Drought could not limit the density of Aedes mosquitos in Thailand. Systematic vector control and vector surveillance programs by public health organizations, if practical, should be continuously conducted to reduce or prevent dengue risk.     

Limited potential for transmission of live dengue virus vaccine candidates by Aedes aegypti and Aedes albopictus

To evaluate the transmission risk of four live dengue (DEN) vaccine candidates developed by the U.S. Army (DEN-1, 45AZ5 PDK 20; DEN-2, S16803 PDK 50; DEN-3, CH53489 PDK 20; and DEN-4, 341750 PDK 20), we tested 3,010 Aedes aegypti and 1,576 Aedes albopictus mosquitoes blood-fed on 21 volunteers who had been administered one of the four vaccine candidates or the licensed yellow fever (YF) vaccine (17D). We used an indirect immunofluorescence assay (IFA) to detect DEN or YF viral antigen in the heads of mosquitoes. Corresponding to the lack of a detectable viremia among volunteers inoculated 8-13 days previously with live DEN-1 or DEN-2 vaccine candidates, only six mosquitoes developed disseminated infections after feeding on these volunteers. These six mosquitoes included 4 of 247 Ae. albopictus fed on volunteers inoculated with the DEN-1 vaccine candidate and 2 of 528 Ae. aegypti fed on volunteers inoculated with the DEN-2 vaccine candidate. Infection was confirmed in each of these IFA-positive mosquitoes by isolating infectious virus from the mosquito's body in Vero-cell culture. None of the 1,252 or the 969 mosquitoes fed on DEN-3 or DEN-4 recipients, respectively, were infected. Overall, dissemination rates in Ae. albopictus and Ae. aegypti were low. Dissemination rates were 0.5%, 0.3%, < 0.1%, and < 0.1% for the DEN-1 through DEN-4 vaccine candidates, respectively. Because of the observed low dissemination rates, it is unlikely that these vaccine viruses would be transmitted under natural conditions.     

Size alters susceptibility of vectors to dengue virus infection and dissemination

The size of arthropod vectors may affect their ability to transmit pathogens. Here we test the hypothesis that body size alters the susceptibility of Aedes aegypti and Aedes albopictus mosquitoes to dengue virus (DENV) infection and subsequent dissemination throughout the body of the mosquito. After feeding on blood containing known quantities of virus, smaller-sized females were significantly more likely to become infected and to disseminate virus than larger individuals. The effects of size were stronger for Ae. aegypti and independent of rearing conditions. Ae. albopictus was more susceptible to DENV infection and had higher virus titer in the body than Ae. aegypti, yet infected Ae. aegypti disseminated DENV more readily than infected Ae. albopictus. These results are consistent with the concept that Ae. aegypti is a more competent vector of DENV and emphasize the importance of body size in determining adult infection parameters.     

Introduction of Aedes albopictus in Gabon: what consequences for dengue and chikungunya transmission?

The 2007 outbreak of chikungunya in Gabon has indicated the potential of this disease to spread beyond its usual range ensuing from the expansion of the mosquito Aedes albopictus . A few cases of dengue (DEN) infection were also detected. Because little is known about the potential for Gabonese mosquito species to transmit both chikungunya and DEN viruses (DENV), we conducted studies to determine the susceptibility of Ae. albopictus and Aedes aegypti collected in Libreville to both viruses by experimental infections. Disseminated infection rates were high for Ae. albopictus infected with chikungunya virus (CHIKV) (66.7–86%) and low with DENV (13–21.4%). Moreover, Ae. aegypti sp. formosus was a less efficient vector of CHIKV than Ae. albopictus . The recent introduction and dissemination of chikungunya associated with the invasion of Ae. albopictus in Africa illustrates the potential for CHIKV to spread to other parts of the world.     

Aedes aegypti in French Guiana: susceptibility to a dengue virus

Twenty-seven samples of Aedes aegypti (F1 generation) from French Guiana were tested for their susceptibility to dengue serotype 2 virus. Very high infection rates were observed by indirect fluorescent antibody (IFA) test. Ae. aegypti samples were pooled according to two groups: the first group (N=10) represented mosquitoes from the urbanized area of Cayenne and surroundings, and the second group (N=17) corresponded to mosquitoes collected in the countryside. Infection rates were found to be similar in these two cases. These findings are discussed in relation with the history of Ae. aegypti in this part of the world.     

Sexual transmission of dengue viruses by Aedes albopictus

Male Aedes albopictus experimentally infected with dengue virus types 1, 2, 3, or 4 transmitted their infection sexually to female Ae. albopictus. Such transmission was enhanced if the females had taken a bloodmeal 2 to 7 days prior to mating. Male Ae. albopictus also transmitted dengue virus vertically to their F1 progeny. Infected progeny were found among those derived from eggs laid greater than or equal to 73 hr after mating but not among those derived from eggs laid prior to that time. This suggests that virus probably was not transmitted directly to ova but, rather, underwent prior replication in the female genital tract. Female Ae. albopictus experimentally infected with dengue type 1 virus did not transmit their infection sexually to males. This finding supports the hypothesis that male mosquitoes naturally infected with dengue virus acquired their infection vertically.     

Potential role of sylvatic and domestic African mosquito species in dengue emergence

Dengue virus 2 (DENV-2) strains that circulate in sylvatic habitats of Senegal and other parts of west Africa are believed to represent ancestral forms that evolved into endemic/epidemic strains that now circulate widely in urban areas of the tropics. Previous studies suggested that the evolution of the endemic/epidemic strains was mediated by adaptation to the peridomestic mosquito vectors Aedes aegypti and Ae. albopictus. We conducted experimental infections using sylvatic and peridomestic Senegalese mosquitoes, and both sylvatic and urban DENV-2 strains to determine if endemic DENV-2 adaptation was vector species specific, and to assess ancestral vector susceptibility. Aedes furcifer and Ae. luteocephalus, probable sylvatic vectors, were highly susceptible to both sylvatic and urban DENV-2 strains. In contrast, sylvatic Ae. vittatus and both sylvatic and peridomestic populations of Ae. aegypti were relative refractory to all DENV-2 strains tested. These results indicate that adaptation of DENV-2 to urban vectors did not result in a loss of infectivity for some African sylvatic vectors. Implications for dengue emergence in west Africa are discussed.