The role of windows of selection and windows of dominance in the evolution of insecticide resistance in human disease vectors

Persistent insecticides sprayed onto house walls, and incorporated into insecticide-treated bednets, provide long-acting, cost-effective control of vector-borne diseases such as malaria and leishmaniasis. The high concentrations that occur immediately post-deployment may kill both

resistant and susceptible insects. However, insecticide concentration, and therefore killing ability, declines in the months after deployment. As concentrations decline, resistant insects start to survive while susceptible insects are still killed. The period of time after deployment, within which mortality of resistant individuals is lower than that of susceptible ones, has been termed the ‘window of selection’ in other contexts. It is recognised as driving resistance in bacteria and malaria parasites, both of which are predominantly haploid. We argue that paying more attention to these mortality differences can help understand the evolution of insecticide resistance. Because insects are diploid, resistance encoded by single genes generates heterozygotes. This gives the potential for a narrower 'window of dominance', within the window of selection, where heterozygote mortality is lower than that of susceptible homozygotes. We explore the general properties of windows of selection and dominance in driving resistance. We quantify their likely effect using data from new laboratory experiments and published data from the laboratory and field. These windows can persist months or years after insecticide deployments. Differential mortalities of resistant, susceptible and heterozygous genotypes, after public-health deployments, constitute a major challenge to controlling resistance. Greater attention to mortality differences by genotype would inform strategies to reduce the evolution of resistance to existing and new insecticides.