Tor Almaas

Eleclrophysiological recordings from single olfactory receptor cells were carried out in the male tobacco budworm moth,Heliothis virescens. Recordings were made primarily from the sensilla trichodea type 1, which are located in the characteristic circumferential rows on the antennae. They possess the longest sensilla hairs as revealed by scanning electron microscopy (SEM). The sensory cells of these sensilla responded specifically to pheromones. Only three types of receptor neurons were found, each tuned to one of the female-produced components. The majority (58%) of the neurons were tuned to the major component (Z)-11-hexadecenal (Z11-16:A1). Another large group (27%) responded specifically to stimulation with (Z)-9-tetradecenal (Z9-14: Al). These two compounds are the most important components of the pheromones as judged by their influence on the behavioral responses of the males. The third type of neurons responded specifically to (Z)-11-hexadecen-1-ol (Z11-16: OH), which may act either as a pheromone component or as an interspecific cue. None of the receptor neurons in the long sensilla trichodea responded specifically to the minor aldehyde components of the pheromone, which have subtle effects on behavior. Mixture experiments provided no evidence that minor components influence the receptor responses to the major components. Olfactory sensilla outside the crosswise rows were also characterized morphologically by SEM. Included in these were sensilla of different lengths, corresponding to a classification as s. basiconica and s. trichodea type 2. Electrophysiological recordings from these sensilla showed that they are involved primarily in host odor reception. However, a few of these neurons responded to pheromones.

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.

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.

Knowledge about the neuronal pathways of the taste system is interesting both for studying taste coding and appetitive learning of odours. We here present the morphology of the sensilla styloconica on the proboscis of the moth Heliothis virescens and the projections of the associated receptor neurones in the central nervous system. The morphology of the sensilla was studied by light microscopy and by scanning- and transmission electron microscopy. Each sensillum contains three or four sensory neurones; one mechanosensory and two or three chemosensory. The receptor neurones were stained with neurobiotin tracer combined with avidin-fluorescein conjugate, and the projections were viewed in a confocal laser-scanning microscope. The stained axons entered the suboesophageal ganglion via the maxillary nerves and were divided into two categories based on their projection pattern. Category one projected exclusively ipsilaterally in the dorsal suboesophageal ganglion/tritocerebrum and category two projected bilaterally and more ventrally in the suboesophageal ganglion confined to the anterior surface of the neuropil. The bilateral projecting neurones had one additional branch terminating ipsilaterally in the dorsal suboesophageal ganglion/tritocerebrum. A possible segregation of the two categories of projections as taste and mechanosensory is discussed.