Host recognition by the tobacco hornworm is mediated by a host plant compound.

It is generally believed that animals make decisions about the selection of mates, kin or food on the basis of pre-constructed recognition templates. These templates can be innate or acquired through experience. An example of an acquired template is the feeding preference exhibited by larvae of the moth, Manduca sexta. Naive hatchlings will feed and grow successfully on many different plants or artificial diets, but once they have fed on a natural host they become specialist feeders. Here we show that the induced feeding preference of M. sexta involves the formation of a template to a steroidal glycoside, indioside D, that is present in solanaceous foliage. This compound is both necessary and sufficient to maintain the induced feeding preference. The induction of host plant specificity is at least partly due to a tuning of taste receptors to indioside D. The taste receptors of larvae fed on host plants show an enhanced response to indioside D as compared with other plant compounds tested.

Octopamine mimics the effects of parasitism on the foregut of the tobacco hornworm Manduca sexta.

The parasitic braconid wasp Cotesia congregata lays its eggs inside the body of the larval stage of its host, the moth Manduca sexta. The Cotesia congregata larvae develop within the hemocoel of their host until their third instar, when they emerge and spin cocoons and pupate on the outer surface of the caterpillar. From this time until their death approximately 2 weeks later, the Manduca sexta larvae show striking behavioral changes that include dramatic declines in spontaneous activity and in the time spent feeding. Coincident with these behavioral changes, it is known that octopamine titers in the hemolymph of the host become elevated by approximately 6.5-fold. Octopamine is an important modulator of neural function and behavior in insects, so we examined hosts for neural correlates to the behavioral changes that occur at parasite emergence. We found that, in addition to the changes reported earlier, after parasite emergence (post-emergence), Manduca sexta larvae also showed marked deficits in their ability to ingest food because of a disruption in the function of the frontal ganglion that results in a significant slowing or the absence of peristaltic activity in the foregut. This effect could be produced in unparasitized fifth-instar larvae by application of blood from post-emergence parasitized larvae or of 10(-6)mol l(-1)d,l-octopamine (approximately the level in the hemolymph of post-emergence larvae). In contrast, blood from parasitized larvae before their parasites emerge or from unparasitized fifth-instar larvae typically had no effect on foregut activity. The effects of either post-emergence parasitized blood or 10(-6)mol l(-1) octopamine could be blocked by the octopamine antagonists phentolamine (at 10(-5)mol l(-1)) or mianserin (at 10(-7)mol l(-1)).

Projection pattern of sensory neurons in the central nervous system of a homeotic mutation of the moth Manduca sexta.

Octopod (Octo) is a mutation of the moth Manduca sexta, which transforms the first abdominal segment (A1) in the anterior direction. Mutant animals are characterized by the appearance of homeotic thoracic-like legs on A1. We exploited this mutation to determine what rules might be used in specifying the fates of sensory neurons located on the body surface of larval Manduca. Mechanical stimulation of homeotic leg sensilla did not cause reflexive movements of the homeotic legs, but elicited responses similar to those observed following stimulation of ventral A1 body wall hairs. Intracellular recordings demonstrated that several of the motoneurons in the A1 ganglion received inputs from the homeotic sensory hairs. The responses of these motoneurons to stimulation of homeotic sensilla resembled their responses to stimulation of ventral body wall sensilla. Cobalt fills revealed that the mutation transformed the segmental projection pattern of only the sensory neurons located on the ventral surface of A1, resulting in a greater number with intersegmental projection patterns typical of sensory neurons found on the thoracic body wall. Many of the sensory neurons on the homeotic legs had intersegmental projection patterns typical of abdominal sensory neurons: an anteriorly directed projection terminating in the third thoracic ganglion (T3). Once this projection reached T3, however, it mimicked the projections of the thoracic leg sensory neurons. These results demonstrate that the same rules are not used in the establishment of the intersegmental and leg-specific projection patterns. Segmental identity influences the intersegmental projection pattern of the sensory neurons of Manduca, whereas the leg-specific projections are consistent with a role for positional information in determining their pattern.

Octopod, a homeotic mutation of the moth Manduca sexta, affects development of both mesodermal and ectodermal structures.

Several aspects of leg development in the moth Manduca sexta were examined using the homeotic mutation Octopod (Octo). This mutation causes a transformation of the ventral epidermis of the first abdominal segment (A1) to that of the third thoracic segment (T3), resulting in the presence of thoracic-like legs on A1. The degree of transformation of A1 is variable, ranging from bumps on the cuticle to fully segmented thoracic-like legs. In the normal thoracic legs, clusters of undifferentiated cells known as differentiation centers are located around the coxal-trochanteral, femoral-tibial, and tibial-tarsal joints. The adult thoracic legs develop from the differentiation centers at metamorphosis. The homeotic legs of the Octopod larvae also have differentiation centers at comparable positions in the homeotic leg. As a result, the number of leg segments in a mutant adult is correlated with the number of segments and differentiation centers that animal had in its larval homeotic leg. Our data suggest that the differentiation center located at the coxal-trochanteral joint forms the adult coxa and trochanter, the center at the larval femoral-tibial joint the adult femur and tibia, and the differentiation center at the larval tibial-tarsal joint the adult tarsus. Homeotic larval legs which include at least a femur have supernumerary muscles, while adult homeotic legs rarely show discrete muscle. The homeotic larval muscles appear to have thoracic identities, based on their attachment points and the timing of their degeneration at the larval-pupal transition. They are innervated by a motoneuron that is normally present in A1 where it innervates the ventral lateral external muscle (VLE). In mutant animals, the same motoneuron innervates all of the homeotic muscles and the VLE. We consider possible mechanisms underlying the development of homeotic muscles and their innervation. At the larval-pupal transition, the VLE in mutant animals degenerates at its normal time, which is 3 days after the degeneration of the homeotic muscles. Thus, despite their common innervation, the two muscle types degenerate according to their own schedules, indicating that the developmental fates of the muscles are not dictated by their innervating neuron but are intrinsic to the muscles themselves.

Multisegmental analyses of acoustic startle in the flying cricket (Teleogryllus oceanicus): kinematics and electromyography.

Tethered, flying Australian field crickets (Teleogryllus oceanicus) stimulated with ultrasound respond with a rapid, short-latency turn from the sound source. We analyzed the kinematics of two behavioral components of this acoustic startle response and recorded electromyograms from the muscles involved in producing them. The two behavior patterns studied were the swing of the metathoracic leg, which has been shown to elicit a short-latency turn, and a lateral swing of the antennae, for which a direct role in steering has not been demonstrated. The kinematic data showed that when a pulse of ultrasound was presented to one side of the animal (1) the contralateral metathoracic leg abducted and elevated, while the ipsilateral leg remained in place, (2) both antennae swung laterally, but the contralateral antenna moved farther than the ipsilateral antenna, (3) increases in stimulus intensity elicited larger movements of the leg and contralateral antenna, while the ipsilateral antenna showed little sensitivity to stimulus intensity, and (4) for the leg, the latency to the onset of the swing decreased and the duration of the movement increased with increasing stimulus intensity. Electromyograms were recorded from the leg abductor M126 and two antennal muscles: the medial scapo-pedicellar muscle M6 and the lateral scapo-pedicellar muscle M7. M7 moves the antenna laterally, M6 moves it medially. Upon stimulation with ultrasound (1) both M126 and M7 showed increasing spike activity with increasing intensity of the ultrasound stimulus, (2) M126 showed a decrease in latency to the first spike and an increase in the duration of spike activity with increasing stimulus intensity, (3) latencies for M6 and M7 were not correlated with stimulus intensity, but M7 had significantly shorter latencies than M6 and the contralateral M7 had significantly shorter latencies than the ipsilateral M7, and (4) the ipsilateral M126 spiked in response to ultrasound in 6 of the 10 animals tested. In these cases, however, latency to the first spike was substantially longer, and the spike frequency was lower than for the muscle's response to contralateral stimuli. We attempt to correlate these electromyogram data with the kinematic data and relate them to the relevance of the two behavior patterns to the execution of an escape response.

Developmental attenuation of the pre-ecdysis motor pattern in the tobacco hornworm, Manduca sexta.

At the culmination of each molt, the larval tobacco hornworm exhibits a pre-ecdysis behavior prior to shedding its old cuticle at ecdysis. Both pre-ecdysis and ecdysis behaviors are triggered by the peptide, eclosion hormone (EH). Pre-ecdysis behavior consists of rhythmic abdominal compressions that loosen the old larval cuticle. This behavior is robust at larval molts, but at the larval-pupal molt the only comparable behavior consists of rhythmic dorso-ventral flexions of the anterior body. These flexions appear to be an attenuated version of the larval pre-ecdysis behavior because (1) they show the same EH dependence, and (2) the motor patterns recorded from EH treated, deafferented larval and pupal preparations are similar except that the pupal pattern is much weaker. Both patterns are characterized by rhythmic, synaptically-driven bursts of action potentials in motoneurons MN-2 and MN-3, which occur synchronously in all segments. However, the synaptic drive to the motoneurons and their resultant levels of activity are reduced during the pupal pre-ecdysis motor pattern, especially in posterior abdominal segments. Although the dendritic arbors of both motoneurons regress somewhat during the larval-pupal transformation, this does not appear to be the primary source of diminished synaptic drive because regression is greatest in the segments in which synaptic inputs remain the strongest. The developmental weakening of the pre-ecdysis motor pattern thus may be due to changes at the interneuronal level.

The effects of behaviourally relevant temperatures on mechanosensory neurones of the grasshopper, Schistocerca americana.

Grasshopper mechanosensory hair neurones respond to displacement of their associated hairs in a temperature sensitive manner: comparable increases in the number of spikes per stimulus result from increases in temperature with constant stimulus strengths and from increasing stimulus strengths at constant temperature. It is therefore not obvious that neurones in the CNS which receive inputs from mechanosensory hairs would be able to distinguish between these two parameters. The temperatures which populations of mechanosensory hairs on the thorax, head and tarsus experienced were measured in freely moving animals. Animals in thermally heterogeneous environments spent 90% of the accounted time in locations where thoracic temperatures of 32-44 degrees C were maintained (the behaviourally 'preferred' range). Head temperatures covered a wider range, and tarsal temperatures the widest. Different populations of mechanosensory hair neurones exhibited different sensitivities to temperature. Thoracic hair neurones were significantly more temperature sensitive than one of the two populations of head hairs studied, and tarsal hairs exhibited a pronounced temperature compensation in the behaviourally 'preferred' range. Wind sensitive head hairs, however, showed exceptionally high temperature sensitivities. There is a negative correlation between the temperature sensitivity of a population of mechanosensory hair neurones and the temperature variability to which those neurones are normally exposed. Implications of this correlation for the central interpretation of mechanosensory input are considered.