The antennae (and palps) come in a multitude of shapes (Figure 1A) but nevertheless conform to the same basic principles (Schneider, 1964). The distal segment of the antennae is covered, to various extents with olfactory sensilla, which show a wide variety of shapes and structures (Schneider and Steinbrecht, 1968) (Figures 1B–1F). Irrespective of form, the olfactory sensilla all share

the same function, namely, to encapsulate and protect Sunitinib mouse the sensitive dendrites of the olfactory sensory neurons (OSNs) (Zacharuk, 1980) (Figure 2A). Although fulfilling the same role, the organization of the peripheral olfactory system of insects is quite different from that of mammals (Figure 2B). The insect antennae have presumably evolved from structures that predominantly mediated mechanosensory input. In primitive terrestrial arthropods, the antennae have great flexibility of movement due to the presence of intrinsic musculature, but owing to the small number of sensilla, quite a poor capacity for chemoreception. The sensillum-rich flagellar antennae found in most insects are, however, void of intrinsic muscles, and are in most lineages specialized structures for detecting odor molecules (Schneider, 1964). Exemptions selleck compound are naturally found, such as in the aquatic water scavenger beetles (Coleoptera: Hydrophillidae), whose antennae actually lack an olfactory function altogether and instead serve as “snorkels,” which are

used to refill internal air reservoirs (Schaller, 1926). Whether antennal Liothyronine Sodium architecture is shaped by the evolutionary necessity to detect certain odor molecules is uncertain. Most likely, the variability in antennal shapes (as seen in Figure 1A) reflects constraints imposed by the physical, rather than the chemical environment of the insects. For example, the delicate plumose antennae of the volant Nevada buck moth in Figure 1A has very likely evolved to capture volatile molecules with high efficiency in air, but would be ill suited to fulfilling the same function for a ground- or soil-dwelling insect. As to why insects are equipped with a second nose, i.e., the maxillary and/or the labial palps, remains unclear.

In several insect species, including the hawk moth Manduca sexta (Lepidoptera: Sphingidae) and the African malaria mosquito Anopheles gambiae (Diptera: Culicoidae), these organs serve a distinct function as they house OSNs detecting CO2, which in both species is a crucial sensory cue for locating resources ( Thom et al., 2004 and Lu et al., 2007). However, in the vinegar fly Drosophila melanogaster (Diptera: Drosophilidae), CO2 detection is accomplished via OSNs on the antennae, and the palp’s OSNs show overlapping response spectra with those of the antennae ( de Bruyne et al., 1999). In the vinegar fly, the palps have instead been suggested to play a role in taste enhancement ( Shiraiwa, 2008). How general such a function would be across insects remains to be investigated.