No abstract

The macroglomerular complex in the primary olfactory center of male moths receives information from numerous pheromone-detecting receptor neurons housed in specific sensilla located on the antennae. We investigated the functional organization of the three glomeruli constituting this complex in Helicoverpa assulta, a unique species among heliothine moths as concerns the composition of the pheromone blend. By tip recordings from the male-specific receptor neurons combined with cobalt-lysine stainings, the axon terminals in the brain were traced and subsequently reconstructed by camera lucida drawings. Some were also reconstructed in a digital form. The results showed that the sensilla could be classified into two functional types. A major category housed two colocalized receptor neurons, one responding to the primary pheromone component cis-9-hexadecenal and the other to the behavioral antagonists cis-9-tetradecenal and cis-9-hexadecenol. Cobalt-lysine applied to this sensillum type consistently resulted in two stained axons, each terminating in one of the two large subunits of the macroglomerular complex: the cumulus or the dorsomedial glomerulus. The second, less frequently appearing sensillum type contained a receptor neuron responding to the second pheromone component, cis-11-hexadecenal. Dye applied to this type resulted in stained axon projections in the ventral glomerulus. In an evolutionary context it is particularly interesting that differences of related heliothine species are reflected in the functional organization of the MGC compartments.

Partial electroantennograms (EAGs) and single cell recordings fromHeliothis virescens males have demonstrated the presence of pheromones receptor neurons in sensilla trichodea type 2 as well as in type 1. This is supported by cobalt tracing experiments, showing that primary axons from the distal flagellum, containing only s. trichodea type 2, project into the macrogiomerulus complex in the male antennal lobes. Four types of finely tuned pheromone receptor neurons were found in males, whereas in females the corresponding neurons responded mainly to host odors. In males the majority (75 and 18%, respectively) were tuned to the majorHeliothis virescens pheromone components (Z)-11-hexadecenal (Z11-16∶A1) and (Z)-9-tetradecenal (Z9-14∶A1). The others (5 and 2%, respectively) responded specifically to (Z)-1 1-hexadecen-1-ol (Z1 1-16∶OH) and (Z)-1 1-hexadecen-1-ol acetate (Z1 1-16∶Ac). No neurons responding selectively to the minor pheromone components were found. The Z11-16∶A1 neurons of both sensilla types possessed similar specificity. However, the sensitivity decreased toward the medial and distal part of the flagellum, where s. trichodea type 2 are located. This suggests that the pheromone concentrations can be detected peripherally by a spatial as well as a temporal mechanism. Differences in temporal response patterns (pronounced phasic vs. tonic component) were found within the same type of neurons, suggesting different ability to encode intermittency of the pheromone plume as well as to mediate maintenance of flight.

Sucrose stimulation of gustatory receptor neurons on the antennae, the tarsi, and the mouthparts elicits the proboscis extension reflex in many insect species, including lepidopterans. The sensory pathways involved in this reflex have only partly been investigated, and in hymenopterans only. The present paper concerns the pathways of the gustatory receptor neurons on the antennae and on the proboscis involved in the proboscis extension reflex in the moth Heliothis virescens (Lepidoptera; Noctuidae). Fluorescent dyes were applied to the contact chemosensilla, sensilla chaetica on the antennae, and sensilla styloconica on the proboscis, permitting tracing of the axons of the gustatory receptor neurons in the central nervous system. The stained axons showed projections from the two appendages in two closely located but distinct areas in the suboesophageal ganglion (SOG)/tritocerebrum. The projections of the antennal gustatory receptor neurons were located posterior-laterally to those from the proboscis. Electrophysiological recordings from the receptor neurons in s. chaetica during mechanical and chemical stimulation were performed, showing responses of one mechanosensory and of several gustatory receptor neurons. Separate neurons showed excitatory responses to sucrose and sinigrin. The effect of these two tastants on the proboscis extension reflex was tested by repeated stimulations with solutions of the two compounds. Whereas sucrose elicited extension in 100% of the individuals in all repetitions, sinigrin elicited extension in fewer individuals, a number that decreased with repeated stimulation.

Discrimination of edible and noxious food is crucial for survival in all organisms. We have studied the physiology of the gustatory receptor neurons (GRNs) in contact chemosensilla (insect gustatory organs) located on the antennae of the moth Heliothis virescens, emphasizing putative phagostimulants and deterrents. Sucrose and the 2 bitter substances quinine and sinigrin elicited responses in a larger proportion of GRNs than inositol, KCl, NaCl, and ethanol, and the firing thresholds were lowest for sucrose and quinine. Variations in GRN composition in individual sensilla occurred without any specific patterns to indicate specific sensillum types. Separate neurons showed excitatory responses to sucrose and the 2 bitter substances quinine and sinigrin, implying that the moth might be able to discriminate bitter substances in addition to separating phagostimulants and deterrents. Besides being detected by separate receptors on the moth antennae, the bitter tastants were shown to have an inhibitory effect on phagostimulatory GRNs. Sucrose was highly appetitive in behavioral studies of proboscis extension, whereas quinine had a nonappetitive effect in the moths.