, 2007). However, by extending the analysis GSK1349572 in vitro to all 12 sequenced Drosophila species, Gardiner and colleagues (2008) found that the proportion of pseudogenized genes did not differ between the specialist and generalist taxa, whereas the endemic species showed significantly more losses than

the mainland species. In their view, small effective population size and genetic drift may rather account for OR gene loss than ecological specialization. Firmly categorizing these species in terms of ecology and demography is however difficult. For example, although D. erecta is specialized upon fruit from Pandanus spp. screwpines, this resource is not continuously available in the habitat. Accordingly, this species must also utilize other resources. Moreover, D. erecta has a restricted and patchy distribution and may thus in fact have a small Volasertib cell line effective population size ( Lachaise et al., 1988). Consequently, examining OR repertoires of additional drosophilid taxa is undoubtedly necessary before any firm conclusions can be drawn. In short, the molecular basis of insect olfaction shows a number of unique features and is characterized by two large gene families, the OBPs and the ORs, which are presumably exclusive to this group of animals. When these two gene families first appear in the insect lineage and whether the initial conquest of land or the diversification

of land plants drove their evolution remains to be determined. All insect genomes to date stem from derived orders. Deep sequencing of species from basal insect orders, as well as from allied hexapod

orders is thus needed in order to understand the evolutionary history of these gene families. Insects have to detect specific volatile information crotamiton in a very complicated chemical environment. How is this feat accomplished? In the vinegar fly and the African malaria mosquito, more or less the complete OR repertoires have been deorphaned, i.e., their key odorant stimuli have been identified. In both species, the ORs display a varying degree of specificity, with certain receptors showing a high degree of selectivity, while others respond to a broad spectrum of compounds (Carey et al., 2010 and Hallem and Carlson, 2006). Response profiles of OSNs, obtained through single sensillum recordings (SSRs) from numerous other insects also suggest a spectrum of OR binding affinities. Perhaps the most well-known specialist OSNs are those detecting pheromones, where OSNs capable of separating two enantiomers with a specificity spanning over more than four decadic concentration steps have been found (Wojtasek et al., 1998). Highly specialized OSNs tuned to host volatiles have been identified from a number of insect species (e.g., Mustaparta et al., 1979, Todd and Baker, 1993 and Tanaka et al., 2009).