Unexpected behavioural adaptation of yellow fever mosquitoes in response to high temperatures.

Temperature is a major ecological driver of mosquito-borne diseases as it influences the life-history of both the mosquito and the pathogen harboured within it. Understanding the mosquitoes' thermal biology is essential to inform risk prediction models of such diseases. Mosquitoes can respond to temperatures by microhabitat selection through thermal preference. However, it has not yet been considered that mosquitoes are likely to adapt to changing temperatures, for example during climate change, and alter their preference over evolutionary time. We investigated this by rearing six cohorts of the yellow fever mosquito Aedes aegypti at two temperatures (24 °C, 30 °C) for 20 generations and used these cohorts to explicitly separate the effects of long-term evolution and within-generation acclimation on their thermal preferences in a thermal gradient of 20-35 °C. We found that warm-evolved mosquitoes spent 31.5% less time at high temperatures, which affects their efficiency as a vector. This study reveals the complex interplay of experimental evolution, rearing temperatures, and thermal preference in Ae. aegypti mosquitoes. It highlights the significance of incorporating mosquito microhabitat selection in disease transmission models, especially in the context of climate change.

Thermal preference of Culicoides biting midges in laboratory and semi-field settings.

Biting midges of the genus Culicoides (Diptera: Ceratopogonidae) are hematophagous insects, and some species can transmit a plethora of pathogens, e.g., bluetongue virus and African horse sickness virus, that mainly affect animals. The transmission of vector-borne pathogens is strongly temperature dependent, and recent studies pointed to the importance of including microclimatic data when modelling disease spread. However, little is known about the preferred temperature of biting midges. The present study addressed the thermal selection of field-caught Culicoides with two experiments. In a laboratory setup, sugar-fed or blood-fed midges were video tracked for 15 min while moving inside a 60 × 30 × 4 cm setup with a 15-25 °C temperature gradient. Culicoides spent over double the time in the coldest zone of the setup compared to the warmest one. This cold selection was significantly stronger for sugar-fed individuals. Calculated preferred temperatures were 18.3 °C and 18.9 °C for sugar-fed and blood-fed Culicoides, respectively. The effect of temperature on walking speed was significant but weak, indicating that their skewed distribution results from preference and not cold trapping. A second experiment consisted of a two-way-choice-setup, performed in a 90 × 45 × 45 cm net cage, placed outdoors in a sheltered environment. Two UV LED CDC traps were placed inside the setup, and a mean temperature difference of 2.2 °C was created between the two traps. Hundred-fifty Culicoides were released per experiment. Recapture rates were negatively correlated with ambient temperature and were on average three times higher in the cooled trap. The higher prevalence of biting midges in cooler environments influences fitness and ability to transmit pathogens and should be considered in models that predict Culicoides disease transmission.

The Phlebotomine sand fly fauna of Switzerland revisited.

Sand flies (Diptera: Psychodidae, Phlebotominae; Newstead, 1911) are widespread in Europe, being particularly common in the Mediterranean region but rare north of the Alps. Thus, Switzerland is an opportune place to investigate the sand fly fauna on both sides of the Alpine crest, in southern sub-Mediterranean climate and northern oceanic temperate climate. We reinvestigated the Swiss sand fly fauna with the aim to assess changes in composition, altitudinal distribution, abundance and seasonality. Thirty-eight sites were investigated with light traps and/or interception sticky traps in 4 years. Ninety and 380 specimens were caught by light traps and sticky traps, respectively, at 15 collecting sites. Four species were identified. Phlebotomus mascittii (Grassi, 1908), Phlebotomus perniciosus (Newstead, 1911) and Sergentomyia minuta (Rondani, 1843) were confirmed in Ticino, and P. mascittii for the first time in neighbouring Grisons. Also, Phlebotomus neglectus (Tonnoir, 1921) is for the first time reported, though at a very low density compared to P. perniciosus at the same site. Its presence in Ticino supports the northward spread observed in Italy. Sand flies were detected north of the Alps at one site only, endorsing a historical report. Overall, the low density of P. perniciosus and very low density of P. neglectus suggest that canine leishmaniosis may not be an important disease risk in Switzerland.

Environment rather than breed or body site shapes the skin bacterial community of healthy sheep as revealed by metabarcoding.

BACKGROUND: The skin is inhabited by a variety of micro-organisms, with bacteria representing the predominant taxon of the skin microbiome. In sheep, the skin bacterial community of healthy animals has been addressed in few studies, only with culture-based methods or sequencing of cloned amplicons. OBJECTIVES: The objectives of this study were to determine the sheep skin bacterial community composition by using metabarcoding for a detailed characterisation and to determine the effect of body part, breed and environment. MATERIALS AND METHODS: Overall, 267 samples were taken from 89 adult female sheep, belonging to three different breeds and kept on nine different farms in Switzerland. From every individual, one sample each was taken from belly, left ear and left leg and metabarcoding of the 16S rRNA V3-V4 hypervariable region was performed. RESULTS: The main phyla identified were Actinobacteriota, Firmicutes, Proteobacteria and Bacteriodota. The alpha diversity as determined by Shannon's diversity index was significantly different between sheep from different farms. Beta diversity analysis by principal coordinate analysis (PCoA) showed clustering of the samples by farm and body site, while breed had only a marginal influence. A sparse partial least squares discriminant analysis (sPLS-DA) revealed seven main groups of operational taxonomic units (OTUs) of which groups of OTUs were specific for some farms. CONCLUSIONS AND CLINICAL RELEVANCE: These findings indicate that environment has a larger influence on skin microbial variability than breed, although the sampled breeds, the most abundant ones in Switzerland, are phenotypically similar. Future studies on the sheep skin microbiome may lead to novel insights in skin diseases and prevention.

Effects of host blood on mosquito reproduction.

Female mosquitoes require blood from their host for egg development. However, the relationship between the composition of host blood and mosquito reproduction, and whether and how this is linked to host selection, remain unclear. A better understanding of these issues is beneficial for mass-rearing of mosquitoes for vector control. This review provides an overview of the currently known effects of blood constituents on mosquito reproduction. Furthermore, it highlights knowledge gaps and proposes new avenues for investigation. We recommend that research efforts be focused on physiological differences between generalist and specialist mosquito species as models to investigate if and how host preference correlates with reproductive output.

Temperature preference of sugar- or blood-fed Aedes japonicus mosquitoes under semi-natural conditions.

Mosquito-borne diseases pose a major burden on humans and animals. Temperature strongly influences the physiology and life cycle of mosquitoes and also the pathogens they transmit. Thermoregulatory behaviour of mosquitoes has been addressed in a few laboratory studies. Here, we expand such studies by investigating the thermal preference when resting of Aedes japonicus, an invasive and putative vector species of many pathogens, in a semi-field setup during summers in a temperate climate. Blood-fed or sugar-fed Ae. japonicus females were released in the late afternoon in a large outdoor cage containing three resting boxes. The next morning, temperature treatments were applied to the boxes, creating a "cool" (over all experiments around 18 °C), and a "warm" (around 35 °C) microhabitat in addition to an untreated "ambient" (around 26 °C) one. The mosquitoes resting within the three boxes were counted five times, every 2 h between 9h and 17h. The highest proportions of mosquitoes (e.g. up to 21% of blood-fed ones) were found in the cool box while both blood-fed and sugar-fed mosquitoes avoided the warm box. The mean resting temperatures of Ae. japonicus were below the ambient temperatures measured by a nearby meteorological station, and this was more pronounced at higher outdoor temperatures and in blood-fed as compared to sugar-fed mosquitoes. Thus, over all experiments with blood-fed mosquitoes, the calculated average resting temperature was 4 °C below the outdoor temperature. As mosquitoes prefer cooler resting places than temperatures measured by weather stations in summer, models to predict mosquito-borne disease outbreaks need to account for the thermoregulatory behaviour of mosquitoes, especially in the wake of climate change.

Thermal preference of adult mosquitoes (Culicidae) and biting midges (Ceratopogonidae) at different altitudes in Switzerland.

Mosquitoes (Diptera: Culicidae) and biting midges (Diptera: Ceratopogonidae) are among the most important vectors of human and veterinary pathogens. For modelling the distribution of these pathogens, entomological aspects are essential, which in turn are highly dependent on environmental factors, such as temperature. In this study, mosquitoes and biting midges were sampled in multiple microclimates at two low (360, 480 meters above sea level, m.a.s.l.) and two high (1250, 1530 m.a.s.l.) altitude locations in Switzerland. Sets of various traps (CO2 -baited CDC, LED-UV, resting boxes, oviposition cups) equipped with dataloggers were placed in transects at five sites with similar vegetation at each location. Only the CDC and the LED-UV traps collected enough insects for analyses. Taxonomic diversity was greater for mosquitoes but lower for biting midges at lower altitudes. Both mosquitoes and biting midges had a thermal preference. Culicoides preferred the traps with warmer microclimate, especially at lower altitudes, whereas mosquito preferences depended on the species, but not on altitude. Relative humidity had a significant positive impact on catches of biting midges but not mosquitoes. To obtain better data on thermal preferences of resting and ovipositing vectors in addition to host seeking individuals, new and improved collecting methods are needed.

Sense the moist and feel the heat: mosquito host and oviposition site detection.

Laursen et al. identified coreceptor Ir93a that is required for humidity and thermal sensing in the mosquito species Anopheles gambiae and Aedes aegypti. Behavioural studies with mutant mosquitoes with disrupted Ir93a revealed that they were less attracted to a blood meal source and oviposition site at close distance.

Fermenting molasses and a synthetic odour blend to attract blood-fed Anopheles coluzzii.

Collecting blood-fed mosquitoes to monitor pathogen presence or to gather information on the host blood meal is often challenging. Fermenting molasses can be used to produce carbon dioxide to attract host-seeking mosquitoes, however, earlier work indicated that it may also attract blood-fed mosquitoes in the field. In the current study, these field results were validated in an experimental setting using a large cage setup with Anopheles coluzzii (Diptera, Culicidae). Blood-fed mosquitoes were indeed attracted to fermenting molasses with the highest attraction at 72 hours post feeding, which was used for subsequent experiments. Next, it was tested if fermentation of molasses is required for attraction, and whether it acts as an oviposition attractant, increases egg laying, or increases mosquito survival. The compounds that could be responsible for attraction were identified by combined electrophysiology and chemical analyses and formulated into a synthetic blend. Fermenting molasses attracted blood-fed mosquitoes in the large cage study, while fermenting sugar and non-fermenting molasses did not. The fecundity of blood-fed mosquitoes increased after feeding on fermenting molasses, however, compounds emanating from molasses did not trigger oviposition. The synthetic blend attracted blood-fed mosquitoes and may be used to determine mosquito host selection and for xenomonitoring, as 'flying syringes' to detect non-vector borne pathogens.

Video analysis of the locomotory behaviour of Aedes aegypti and Ae. japonicus mosquitoes under different temperature regimes in a laboratory setting.

Mosquito-borne diseases impose a high burden on human and animal health. Temperature strongly influences the physiology and life cycle of mosquitoes, but also the development and/or propagation of the pathogens they transmit. Thus, the vector capacity of mosquitoes depends strongly on temperature and their behavioural thermoregulation through microhabitat selection. Expanding on a previous study of static thermal preferences, the locomotory dynamics of temperate Aedes japonicus (reared from eggs collected in the field) and tropical Ae. aegypti (from a laboratory colony) was investigated at constant temperatures (10 °C, 25 °C, 40 °C) and in temperature gradients (10-20 °C, 20-30 °C, 30-40 °C). Blood-fed or non-blood-fed female mosquitoes were released in groups of 15 individuals into a Plexiglas box positioned on two thermoregulators connected by an aluminium plate to automatically monitor by video analysis mosquito flying, walking and resting duration, covered distances and velocity. Mosquitoes were predominantly resting, followed by walking and flying. At constant 10 °C, flights were rare and brief, and walking was slow. Most activity was observed at 25 °C for Ae. japonicus and 40 °C for Ae. aegypti. In the 30-40 °C gradient, activity of Ae. aegypti increased towards the cold end, suggesting active avoidance of very high temperatures. In the 20-30 °C gradient, edge effects were prominent, nevertheless revealing a greater proportion of mosquitoes gathered at the cooler end. Video analysis showed that this effect was not caused by a cold trap but represents true thermal preference. In the coolest gradient (10-20 °C), mosquitoes were active in all sectors without displaying a preference for either side. Overall, both the tropical and temperate mosquito species preferred cooler temperatures and actively avoided the hottest temperatures. Further studies with infected mosquitoes should provide important insights for developing models of vector-borne disease outbreaks.

Xenosurveillance proof-of-principle: Detection of Toxoplasma gondii and SARS-CoV-2 antibodies in mosquito blood meals by (pan)-specific ELISAs.

Blood-fed insects can be used to analyse the host blood for circulating vertebrate pathogens or antibodies directed against them. We tested whether naturally acquired antibodies in different host species can be detected by host-specific and pan-specific ELISAs in mosquito blood meals. Cat- and alpaca-specific ELISAs could detect antibodies against Toxoplasma gondii or SARS-CoV-2 in blood meals of Aedes japonicus for 48 and at least 24 h, respectively. In the pan-specific ELISA, a conjugated protein A/G and anti-IgY were used to detect antibodies of mammalian and bird hosts. Thus, Toxoplasma antibodies could be detected in mosquitoes fed on blood from humans, chicken, pig, and sheep up to 72 h after the blood meal. The results, however, demonstrated differences in sensitivities between different host species, and the assay requires further evaluation. Xenosurveillance with antibody detection in mosquito blood meals can be an additional surveillance tool that would especially be helpful when it is difficult to sample the potential animal reservoirs.

Skin bacterial volatiles: propelling the future of vector control.

The skin microbiota plays an essential role in the protection against pathogens. It is our skin microbiota that makes us smell different from each other, rendering us more or less attractive to mosquitoes. Mosquitoes exploit skin bacterial odours to locate their hosts and are vectors of pathogens that can cause severe diseases such as malaria and dengue fever. A novel solution for long-lasting protection against insect vectors of disease could be attained by manipulating the bacterial commensals on human skin. The current options for protection against biting insects usually require topical application of repellents that evaporate within hours. We discuss possible routes for the use of commensal bacteria to create a microbial-based repellent.

Potential mechanical transmission of Lumpy skin disease virus (LSDV) by the stable fly (Stomoxys calcitrans) through regurgitation and defecation.

Lumpy skin disease (LSD) is a viral disorder of cattle caused by the lumpy skin disease virus (LSDV) which can induce severe infections leading to high economic losses. Being of African origin, the first LSD outbreaks in Europe occurred in Greece and later in the Balkan region. Little is known about the mode of transmission, especially in relation to the potential role of arthropods vectors. The purpose of our study was to investigate the role of Stomoxys calcitrans in the transmission of LSDV and their presence at different farms in Switzerland. Laboratory-reared flies were exposed to LSDV spiked-blood and incubated under a realistic fluctuating temperature regime. Body parts, regurgitated blood, and faecal samples were analysed by qPCR for the presence of viral DNA and infectious virus at different time points post-feeding (p.f.). LSDV DNA was detected in heads, bodies, and regurgitated blood up to three days p.f. and up to two days p.f. in the faeces. Infectious virus was isolated from bodies and faeces up to two days and in the regurgitated blood up to 12 h p.f. There was no increase in viral load, consolidating the role of S. calcitrans as mechanical vectors for LSDV. Stomoxys flies were present at all eight farms investigated, including a farm located at 2128 m asl. The persistence of LSDV in S. calcitrans in combination with the long flight ranges of this abundant and widespread fly might have implications on LSD epidemiology and on implementing control measures during disease outbreaks.

Spatial repellency and vapour toxicity of transfluthrin against the biting midges Culicoides nubeculosus and C. sonorensis (Ceratopogonidae).

Biting midges (Diptera; Ceratopogonidae; Culicoides spp.) are biological vectors of disease agents, and they cause nuisance and insect bite hypersensitivity. Currently there are no effective means to control biting midges as screening is impractical and the application of insecticides or repellents is of limited efficacy. Spatial repellents have the advantage over contact repellents that they can create a vector-free environment. Studies have shown the efficacy of spatial repellents to protect humans against mosquitoes, also outdoors, but no data are available for biting midges. We tested the spatial repellency and toxicity (knockdown effect) of the volatile pyrethroid transfluthrin against the laboratory-reared biting midges Culicoides nubeculosus (Meigen) and Culicoides sonorensis (Wirth and Jones) and the mosquito Aedes aegypti (Linnaeus) in a high-throughput tube setup. Observations were made 15, 30 and 60 min. after application of the repellent. In addition to transfluthrin, the non-volatile pyrethroid permethrin and DEET, the gold standard of repellents, were included. Spatial repellency by transfluthrin was observed against both biting midge species and Ae. aegypti, already at the first observation after 15 min. and at much lower concentrations than DEET. Permethrin was spatially repellent only to C. sonorensis at the highest concentration tested (10 µg/cm2). Knockdown of biting midges and mosquitoes by transfluthrin, both by vapour or contact toxicity, was observed even at low concentrations. DEET had little to no effect on the knockdown of the insects, neither by direct contact nor vapour toxicity, while permethrin caused a high proportion of knockdown when direct contact was possible. In case these results can be confirmed in field experiments, spatial repellents could become a novel tool in integrated control programmes to reduce biting by Culicoides spp.

Thermal preferences of subtropical Aedes aegypti and temperate Ae. japonicus mosquitoes.

Temperature is an important determinant affecting the capacity of disease vectors like mosquitoes (Culicidae) to transmit disease agents. Although the impact of temperature on vector-borne disease dynamics has been studied intensively, the actual temperature encountered by the vector in a heterogeneous landscape is rarely taken into account. If disease vectors have temperature preferences and therefore select specific microhabitats, this would substantially influence key life history traits that determine transmission intensity. The thermal preferences of subtropical Aedes aegypti and temperate Ae. japonicus mosquitoes were investigated in a temperature gradient set-up consisting of a Plexiglas box on top of an aluminium plate on two thermal regulators. Blood-fed (one day after feeding) and unfed (non-blood-fed) mosquitoes were released in small (15-20 °C, 20-25 °C, 25-30 °C) and large (15-30 °C, 30-45 °C) temperature gradients to assess their thermal preferences after 15 min. Additionally, the effect of humidity was investigated in a two-choice chamber setup. Both mosquito species avoided higher temperatures, pronouncedly dangerously high temperatures in the 30-45 °C gradient. At lower temperatures, blood-fed mosquitoes preferred the cooler sides of the 20-25 °C and 25-30 °C gradient, which were all below their rearing temperature. In the lowest gradient of 15-20 °C, no preferences were found. The thermal preference of unfed mosquitoes was similar to that of the blood-fed mosquitoes. No humidity preference or effect of humidity on temperature preferences was observed within the tested range (40-90%). The set-up allows for assessing the thermal preference of mosquitoes under controlled conditions. The observed preference of mosquitoes for cooler temperatures would increase their longevity and slow down pathogen development. If similar microhabitat selection is observed in the field, vector borne disease models should be adjusted accordingly.

Optimisation and field validation of odour-baited traps for surveillance of Aedes aegypti adults in Paramaribo, Suriname.

BACKGROUND: Emerging arboviral diseases like Zika, dengue and chikungunya that are transmitted by Aedes aegypti mosquitoes, are increasingly threatening human health. Blends of human-like synthetic chemical attractants can be used to attract host-seeking mosquitoes. The aim of this study was to test new combinations of traps and odour baits in the laboratory, followed by testing the best candidates in the field to improve Ae. aegypti monitoring and surveillance. METHODS: First, the BG-Suna trap was evaluated for capturing laboratory-reared Ae. aegypti by testing normal and inverted positions in screen cage tests. Secondly, the attractiveness of the MB5 blend, CO2, and their combination was tested. Thirdly, we tested the attractiveness of different trap types (BG-Suna, BG-Sentinel, MM-X and CDC light trap). Finally, we confirmed laboratory results in the field in Paramaribo, Suriname, using the MB5 and BG-Lure odour blends, CO2 and the BG-Sentinel and BG-Bowl trap using a Latin Square design. RESULTS: The MB5 blend in combination with CO2 outperformed traps baited only with CO2 or MB5 in screen cage tests (P < 0.0001). The BG-Sentinel trap performed equally well as the inverted BG-Suna and was taken to the field (P = 0.729). In the field, we captured Ae. aegypti, Cx. quinquefasciatus and Cx. nigripalpus. We confirmed the laboratory results and found that the combination of the MB5 blend and CO2 almost doubled Ae. aegypti female captures (P = 0.004) and more than doubled Culex spp. female captures (P = 0.005) compared to using only CO2. Interestingly, the MB5 blend outperformed the commercially available BG-Lure, in the BG-Sentinel (P < 0.001). The BG-Bowl also attracted Ae. aegypti when baited with the MB5 blend in similar numbers as the BG-Sentinel baited with the MB5 (P = 0.362). CONCLUSIONS: Our study demonstrated that the BG-Sentinel trap baited with the MB5 blend and CO2 outperforms the current golden standard (BG-Sentinel trap with BG-Lure) for monitoring Ae. aegypti females and males, in both laboratory and field experiments. The BG-Bowl baited with the MB5 blend is a good candidate for home use. Finally, the results show that CO2 is an indispensable component of the attractive blend.

Effect of overwintering on survival and vector competence of the West Nile virus vector Culex pipiens.

BACKGROUND: West Nile virus (WNV) is a mosquito-borne virus that is mainly transmitted among birds by Culex pipiens mosquitoes. The species Cx. pipiens consists of two biotypes named pipiens and molestus, which together can form hybrids. One of the major distinctions between the biotypes is their overwintering behaviour. Adults of biotype pipiens diapause during winter, whereas biotype molestus remains actively blood-feeding. Diapausing may affect survival and vector competence of biotype pipiens. The aims of this study were therefore to identify the biotype composition of diapausing Cx. pipiens mosquitoes, to quantify survival throughout the autumn and winter months, and to determine effects of overwintering on vector competence of emerging Cx. pipiens mosquitoes for WNV. METHODS: Diapausing mosquitoes were collected at two typical overwintering locations in the Netherlands. A selection of Cx. pipiens mosquitoes was identified to biotype using real-time PCR. Survival of diapausing Cx. pipiens mosquitoes during autumn and winter was monitored by placing cages with either field-collected or laboratory-reared females in houses and sheds. Vector competence of field-collected (diapausing) and laboratory-reared (non-diapausing) Cx. pipiens mosquitoes was determined to gain insight in the effect of overwintering on WNV transmission. RESULTS: The majority (92%) of diapausing Cx. pipiens females were identified as biotype pipiens. More than 70% of diapausing Cx. pipiens mosquitoes was able to survive for more than four months in sheds, whereas diapausing in houses resulted in 100% mortality in that same period. In contrast, non-diapausing Cx. pipiens biotype pipiens mosquitoes reared in the laboratory died within a week in both houses and sheds. Vector competence of Cx. pipiens mosquitoes that had diapaused during the autumn and winter months was comparable to non-diapausing laboratory-reared mosquitoes. CONCLUSIONS: This study confirms that the majority of Cx. pipiens mosquitoes in their typical overwintering site belongs to the pipiens biotype. It shows that more than two-third of diapausing Cx. pipiens mosquitoes is able to survive winter under sheltered winter conditions. Finally, vector competence for WNV of mosquitoes that emerge from overwintering sites is not affected by their relatively old age.

Do apes smell like humans? The role of skin bacteria and volatiles of primates in mosquito host selection.

Anthropophilic mosquitoes are effective vectors of human disease because of their biting preferences. To find their host, these mosquitoes are guided by human odours, primarily produced by human skin bacteria. By analysing the skin bacterial and skin volatile profiles of humans, bonobos, chimpanzees, gorillas, lemurs and cows, we investigated whether primates that are more closely related to humans have a skin bacterial community and odour profile that is similar to that of humans. We then investigated whether this affected discrimination between humans and closely related primates by anthropophilic and zoophilic mosquitoes that search for hosts. Humans had a lower skin bacterial diversity than the other animals and their skin bacterial composition was more similar to that in other primates than it was to the skin bacteria of cows. Like the skin bacterial profiles, the volatile profiles of the animal groups were clearly different from each other. The volatile profiles of cows and lemurs were more closely related to the human profiles than expected. Human volatiles were indeed preferred above cow volatiles by anthropophilic mosquitoes and no preference was observed when tested against non-human primate odour, except for bonobo volatiles, which were preferred over human volatiles. Unravelling the differences between mosquito hosts and their effect on host selection is important for a better understanding of cross-species transmission of vector-borne diseases.

Plasmodium-associated changes in human odor attract mosquitoes.

Malaria parasites (Plasmodium) can change the attractiveness of their vertebrate hosts to Anopheles vectors, leading to a greater number of vector-host contacts and increased transmission. Indeed, naturally Plasmodium-infected children have been shown to attract more mosquitoes than parasite-free children. Here, we demonstrate Plasmodium-induced increases in the attractiveness of skin odor in Kenyan children and reveal quantitative differences in the production of specific odor components in infected vs. parasite-free individuals. We found the aldehydes heptanal, octanal, and nonanal to be produced in greater amounts by infected individuals and detected by mosquito antennae. In behavioral experiments, we demonstrated that these, and other, Plasmodium-induced aldehydes enhanced the attractiveness of a synthetic odor blend mimicking "healthy" human odor. Heptanal alone increased the attractiveness of "parasite-free" natural human odor. Should the increased production of these aldehydes by Plasmodium-infected humans lead to increased mosquito biting in a natural setting, this would likely affect the transmission of malaria.

Mechanisms of Plasmodium-Enhanced Attraction of Mosquito Vectors.

Evidence is accumulating that Plasmodium-infected vertebrates are more attractive to mosquitoes than noninfected hosts, particularly when high levels of gametocytes are present. Changes in host odour have been suggested as a likely target for parasite manipulation because olfactory cues are crucial to mosquitoes in search of a bloodmeal host. This review discusses two routes that may lead to such changes: (i) direct emission of volatile products from malaria parasites, and (ii) changes in skin microbial composition that could lead to changes in the vertebrate odour profile. Here we synthesize what is known and suggest how further research can increase our understanding of the mechanisms of parasite manipulation of host attractiveness.