Organization of a sensory neuropile in the auditory pathway of two groups of Orthoptera.

The anterior intermediate sensory neuropile (aISN) is a prominent neuropile in the ventral nerve cord of locusts and bushcrickets. Previous studies have shown that it receives its main sensory input from auditory receptors. In this paper we examine the structural and physiological relationship between tympanal receptor terminations and the dendrites of sound-sensitive interneurones in the homologous neuropile of locusts and bushcrickets. Each individual receptor fibre of the bushcricket terminates in a somewhat different target area of the neuropile. The ordering is with respect to the characteristic frequency of the fibres (tonotopic) in the anterior-posterior and dorsoventral axis. In the locust, representatives of the four tympanal receptor groups branch in different areas of the aISN. Most of the dorsal neuropilar region, and the anterior ventral region, do not receive input from tympanal receptors. The dendrites of identified sound-sensitive interneurones were examined in the context of this afferent projection. Local interneurones as well as intersegmental interneurones in bushcrickets have dendritic branches in the whole aISN or part of it and thus overlap with at least some receptors. By recording intracellularly from their main neurites, short-latency synaptic potentials were found in response to receptor spikes indicating monosynaptic input. The tuning of these neurones could be predicted by their dendritic morphology. In contrast, in the locust only local and bisegmental neurones are monosynaptically connected with tympanal receptors, but not the studied intersegmental neurones. This is consistent with the finding that most or all branches of intersegmental neurones lie in the dorsal area of neuropile where no receptors terminate. Anatomical and physiological evidence is presented for identified local neurones providing the excitatory and inhibitory synaptic input for such intersegmental neurones. The difference in the basic wiring diagram in the homologous neuropile of the two orthopteran groups is discussed with respect to the possible different roles that sound plays in their behaviour.

Spatial segregation of synaptic inputs and outputs in a locust auditory interneurone.

Using intracellular injection of cobalt, processing for electron microscopy, and using on-section silver intensification, we have investigated the distribution of synapses on the metathoracic arborization of the TN1 intersegmental interneurone in the auditory system of the locust. Previous studies have indicated that inputs to this neurone from tympanal afferents are spatially segregated from its outputs to higher-order inhibitory interneurones. Branches in the area of input are characterised by a smooth appearance in Lucifer Yellow preparations, whereas branches in the main output area appear beaded. Sampling different areas of the TN1 arborization, we have found marked differences in the respective densities of input and output synapses. Labelled profiles in areas of beaded branching have an extremely high density of presynaptic sites, with a much smaller number of synaptic inputs. Smooth branches bear many input synapses, but we have found no output synapses on them. Our results indicate a good correlation between branch beading and presynaptic function and between smooth branches and postsynaptic function. These correlations enable us to predict synapse distribution on other insect neurones with a similar division in branch morphology and also extend our knowledge of the functioning of TN1 itself.