Connectivity pattern of the cercal-to-giant interneuron system of the American cockroach

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Abstract
1. The pattern of connectivity between identified cercal afferents and the three largest giant interneurons (GIs 1, 2, and 3) of the American cockroach was investigated by intracellular methods. These three GIs all have different directional response sensitivities and appear to be especially important in initiating the short latency escape behavior of the American cockroach. 2. One of the interneurons, GI 1 responds to wind from all four quadrants of space about the animal. However, it clearly shows a greater ipsilateral versus contralateral (relative to the GI's axon within the nerve cord) wind sensitivity. In contrast, the directional sensitivity of GI 2 is more nearly bilaterally symmetrical. Both of these interneurons receive excitatory synaptic input from the sensory cells of the nine most prominent columns (a, d, g, f, h, i, k, l, and m) of filiform hairs of the ipsilateral cercus. 3. The nine ipsilateral inputs all made roughly equivalent strength excitatory connections with GI 1. The connectivity pattern to GI 2 was the same as that to GI 1 except that the connection strength for two of the nine columns, h and i, was substantially stronger to GI 2 than to GI 1. The remaining seven sensory columns all make equivalent strength connection with GI 2. 4. Only select columns of contralateral sensory cells made synaptic connection with GIs 1 and 2. All detectable connections produced subthreshold depolarizations. 5. The response curve of GI 3 is more sharply restricted in space than that of either GI 1 or 2 and this interneuron only responds to wind stimuli originating from in front of the animal. GI 3 received excitatory synaptic input only from ipsilateral columns d, f, g, i, and k, all of which have their best excitatory directions well within the boundaries of the response curve of GI 3. Columns a and l with best excitatory directions near the edges of the response curve of GI 3 made no detectable connection. The remaining two columns (h and m) with best excitatory directions well outside the boundaries of the response curve of GI 3 provided inhibitory input. 6. GI 3 received synaptic input from contralateral columns d, f, g, h, i, k, and m.(ABSTRACT TRUNCATED AT 400 WORDS)