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DESCRIPTION: Transmission reduction is a key component Andrea sauerwine fuck nederland global efforts to control and eliminate malaria; yet, it is unclear how the density of transmission stages gametocytes influences infection proportion of mosquitoes infected. Human to mosquito transmission was assessed using direct mosquito feeding assays conducted in Burkina Faso and Kenya.

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Transmission reduction is a key component of global efforts to control and eliminate malaria; yet, it is unclear how the density of transmission stages gametocytes influences infection proportion of mosquitoes infected. Human to mosquito transmission was assessed using direct mosquito feeding assays conducted in Burkina Faso and Kenya.

In a site in Burkina Faso, children harbour more gametocytes than adults though the non-linear relationship between gametocyte density and mosquito infection means that per person they only contribute slightly more to transmission. This method can be used to determine the reservoir of infection in different endemic settings.

Interventions reducing gametocyte density need to be highly effective in order to halt human—mosquito transmission, although their use can be optimised by targeting those contributing the most to transmission. The most severe form is caused by the parasite Plasmodium falciparumwhich can reside within red blood cells and thus evade the human immune system. Plasmodium is transmitted between humans by mosquitoes. When a mosquito takes a blood meal from an individual infected with the parasite, the insect ingests Plasmodium gametocytes i.

Although malaria is both preventable and curable, the mortality rates in many African countries remain high, especially among children. Reducing the transmission of malaria to mosquitoes is one of the primary goals in the global effort to control and eliminate the disease.

While a range of drugs and vaccines that specifically try to reduce transmission are in development, non-medical interventions such as mosquito nets and insecticide spraying can quickly and effectively reduce infection rates. Here, Churcher et al. They found that the transmission occurs readily at very low gametocyte densities. Moreover, the transmission rate remains relatively stable as the density increases, before increasing significantly when the density reaches around cells per microlitre.

By identifying the groups that contribute most Andrea sauerwine fuck nederland transmission, and targeting resources to reduce gametocyte density in those individuals, it could be possible to greatly reduce the number of infected mosquitoes and, therefore, the number of infected humans. The malaria parasite is transmitted among humans by anopheline mosquitoes. Male and female transmission stages gametocytes are ingested by the mosquito and reproduce sexually in its stomach before developing into oocysts.

Once oocysts establish, it is assumed that mosquitoes will form infectious sporozoites, so the proportion of mosquitoes developing oocysts is used as a measure of mosquito infectivity. In the human malaria parasite Plasmodium falciparummosquito infection is thought to increase with the number of gametocytes ingested by the mosquito Jeffery and Eyles, ; Graves et al. However, several studies have failed to find an association Boudin et al.

Unlike conventional microscopy, this technique enables gametocyte densities to be quantified over the entire epidemiologically relevant range. Transmission reduction is now a key component of global efforts to control and eliminate malaria Alonso et al. A wide range of novel transmission-reducing drugs and vaccines are currently under development, which aim to reduce malaria incidence by restricting human to mosquito transmission. The transmission-reducing ability of front-line therapeutics such as artemisinin-based combination therapies [ACTs], or potential combinations of ACTs with gametocytocidal drugs is also gaining increased attention WHO, ; White, It is likely that these interventions will be partially effective, but it remains unclear how much they need to reduce transmission from humans to mosquitoes in a particular location, and to which age groups they should be delivered in order to halt malaria transmission.

Mathematical models of malaria transmission tend not to include explicitly the relationship between gametocytaemia and Andrea sauerwine fuck nederland infectivity, opting, for simplicity, to assume density-independent transmission probabilities Smith et al. Greater complexity, however, will be required to capture fully malaria population dynamics following the introduction of drugs and vaccines that specifically target transmission stages.

More detailed mathematical models will allow the full impact of this change in strategy to be assessed in a range of different endemic settings. The optimal use of transmission-reducing interventions will also require a better understanding of the human reservoir of infection, which is likely to vary between different endemic settings.

Various studies have attempted Andrea sauerwine fuck nederland estimate the relative contribution of different age groups to mosquito infection using skin or membrane feeding assays Muirhead-Thomson, ; Graves et al.

Andrea sauerwine fuck nederland studies are logistically complicated and expensive, so more efficient methods of determining the reservoir of infection are required in order to target interventions optimally. A thorough understanding of the factors determining mosquito infection would allow mathematical models to predict the relative contribution of different groups to transmission from cross-sectional surveys, both before and after Andrea sauerwine fuck nederland introduction of transmission-reducing interventions.

Other covariates that may influence infection such as asexual parasite density measured by microscopy and host age were also included to improve predictions of human to mosquito transmission. These results are combined with data from a cross-sectional survey conducted in a high transmission setting in Burkina Faso to predict the relative contribution of different age groups to overall malaria transmission.

The relationship between the number of gametocytes in the blood and Andrea sauerwine fuck nederland infection was found to be highly non-linear Figure 1A. Plasmodium falciparum infects mosquitoes at the very low gametocyte densities that predominate in natural infections and may not be detected using standard microscopy Bousema and Drakeley, The best-fit model predicts that increasing density from 1 to gametocytes per microlitre does Andrea sauerwine fuck nederland appreciably increase infection.

Gametocyte density on an arithmetic scale was a better predictor of mosquito infection than gametocyte density on a logarithmic scale. Including age improves the fit of the model suggesting that age is an important Andrea sauerwine fuck nederland, although the Bayesian credible intervals include 0.

The complex shape of the relationship between gametocyte density and mosquito infection will influence the success of transmission-reducing interventions and may explain why their ability to reduce the proportion of infected mosquitoes has been shown to depend on the gametocyte density of the host Churcher et al.

The same intervention efficacy at reducing gametocyte density in a host with gametocytes per microlitre would cause an appreciable reduction in human to mosquito transmission from that individual.

Figure 1B can Andrea sauerwine fuck nederland used to estimate how a reduction in the number of gametocytes will equate to a reduction in the proportion of mosquitoes becoming infected, and hence mosquito to human transmission. How this relates to the incidence of malaria and subsequent disease will depend, in part, on the degree of immunity in the human population. In the cross-sectional survey in Burkina Faso, Andrea sauerwine fuck nederland harbour more gametocytes than adults, with 10 year olds having five times as many gametocytes compared to the 50 year olds Figure 2A.

Results indicate that, at the time of the survey conducted in Burkina Faso, the peak in the reservoir of infection occurs at an earlier age than the peak in gametocyte density, which Andrea sauerwine fuck nederland improve the cost effectiveness of school-based programmes.

The complex shape of the relationship Andrea sauerwine fuck nederland gametocyte density and mosquito infection elucidated here will influence our understanding Andrea sauerwine fuck nederland the population dynamics of falciparum malaria and will determine the success of transmission-reducing interventions.

The efficiency with which P. Gametocytes can infect mosquitoes at very low densities, despite the mosquito needing to ingest both male and female parasites in the same blood meal. After this initial increase in infection, considerably more gametocytes are required in order to increase further the proportion of mosquitoes developing oocysts.

The causes of this are unclear, but experimental systems may be informative. Data from the rodent malaria model Plasmodium berghei Sinden et al. Beyond gametocytes per microlitre mosquito infection rises again although QT-NASBA measurement error makes the exact gametocyte density for this transition relatively uncertain.

If an intervention can keep gametocyte density beneath gametocytes per microlitre in a low transmission area, it may be sufficient to push the basic reproduction number of malaria beneath one and eliminate the disease. This means that evaluating interventions solely on their ability to reduce microscopically detectable gametocytes, as has been suggested Graves et al.

Including information on asexual parasite density as measured by microscopy significantly improved the fit of the model, with hosts harbouring intermediate densities having the highest infectivity. It is unclear whether asexual parasites directly influence parasite infectivity or are associated with other here unmeasured variables such as the multiplicity of infection Nsango et al.

Plasmodium falciparum has asynchronous waves of asexual and sexual parasites. Circulating gametocytes take 2—3 days to mature before they can infect mosquitoes Lensen et al.

On average, older children were more infectious, supporting previous indications that transmission-reducing immunity may predominate in young children Drakeley et al. Despite this, they are considerably more accurate than conventional microscopy. Molecular techniques may be highly precise in a state-of-the-art laboratory, but the practicalities of collecting and processing samples in non-ideal field settings increases the risks of incurring measurement error.

Even though uncertainty is generally accepted by those conducting the experiments, measures of precision such as confidence intervals or standard errors Andrea sauerwine fuck nederland such density estimates are rarely reported in the scientific literature.

Methods such as those presented in this article can allow robust quantitative insight to be gained from uncertain molecular methods. Care should be taken when interpreting gametocyte density estimates as it has been suggested that the marker used in this analysis Pfs25 mRNA might be parasite female specific Schneider, Even if the current QT-NASBA method does only detect female gametocytes, it will neither change the qualitative conclusions of this study nor their application if other studies use the same technique.

However, direct comparison of our results with those of studies using different methods for quantifying gametocyte density should be aware that the shape of the relationship may differ. Gametocyte density measurement error is likely to cause the majority of the uncertainty seen in model outputs, although the complexity of the membrane feeding assay is also likely to contribute.

The proportion of mosquitoes infected by Plasmodium -infected blood is known to vary substantially within and between studies. This is Andrea sauerwine fuck nederland part due to methodological issues with the membrane feeding assay Bousema et al. Further standardising the membrane feeding assay, improving the accuracy of QT-NASBA technique and the inclusion of additional covariates such as estimates of gametocyte maturity, which have been investigated in Plasmodium vivax ; Chansamut et al.

It would also permit the patterns described here to be checked for consistency across time and space, and enable a wider range of functional forms to be tested. Care should also be taken when interpreting the results of feeding assays as the mosquitoes used, and their biting behaviour is likely to be different from that seen in wild mosquitoes in field situations Bousema et al.

Transmission reduction will become increasingly important as areas approach local elimination. Identifying host age groups that contribute most to the reservoir of infection will allow the optimal targeting of malaria control. In the high transmission site in Burkina Faso, young children harbour the majority of gametocytes but are only slightly more infective to mosquitoes than adults. The per person contribution of adults estimated here is considerably greater than that predicted by other mathematical models Ross et al.

The contribution of different age groups to overall transmission will depend on local demography, age-dependent protection by malaria interventions such as the use of bed nets or time spent in the house and therefore personally protected by indoor residual sprayingand human to mosquito contact patterns. The last two of these are difficult to measure and poorly understood, reducing our ability to accurately predict the sources of infection.

The reservoir of infection is likely to vary between endemicity settings and over time so multiple cross-sectional surveys may be required. The relationship between gametocyte density and mosquito Andrea sauerwine fuck nederland will allow estimates of the reservoir of infection from cross-sectional gametocytaemia surveys without the need for logistically complicated mosquito feeding assays.

The results of this article should be included within mathematical models that capture changes in gametocyte density with time Lawpoolsri Andrea sauerwine fuck nederland al.

This article provides insights into the relationship between gametocyte density and mosquito infection that are needed to predict the outcome of transmission-reducing interventions. It shows that, given the ability of very low P. The complex and non-linear shape of the relationship between gametocyte density and mosquito infection may explain why the ability of an interventions to reduce the proportion of infected mosquitoes has been shown to depend on the parasite load of the host Churcher et al.

Gametocyte density also changes with host age so the effectiveness of transmission-reducing interventions that target gametocytes will also vary with age. This should be considered in clinical trials of transmission-reducing candidate drugs and vaccines. Like all diagnostic methods, it is prone to measurement error. To understand fully the associated uncertainty, it is important to appreciate the causes of the variability and how the quantification process might magnify uncertainty of point density estimates.

Here, the uncertainty is quantified by repeatedly testing samples with known gametocyte density and fitting a hierarchical mathematical model.

The assay measures time to positivity TTPwhich is the time it takes for the number of target amplicons detected to exceed a defined threshold Schneider et al. The Andrea sauerwine fuck nederland between TTP and gametocyte density is estimated by fitting a linear regression to TTP estimates generated using a sample with known gametocyte density a fold dilution series of in vitro cultured gametocytes ranging from 10 6 to 10 1 gametocytes per millilitre.

Let the observed TTP be denoted by Y then. A full list of the parameters is given Andrea sauerwine fuck nederland Table 1. Here, we adapt this approach for use in a Bayesian hierarchical model. Here, we use the most parsimonious approach that does not require assignment of previous distributions to the unknown gametocyte densities.

Rather, uncertainty in the estimated regression coefficients of equation 1 is propagated numerically via equation 2 to yield uncertainty in the estimated gametocyte densities.

Since all the calibration line data have a relatively steep gradient, the difference between the different methods will be relatively minor, and all will generate sensibly tight confidence interval estimates. Calibration lines can vary between runs and batches of reagents. Here, we define an experiment as being a single plate run with the same batch of reagents each of which will have its own dilution series and samples with unknown density.

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