The code for stimulus direction in a cell assembly in the cockroach

The cockroach Periplaneta americana responds to the approach of a predator by turning away. A gentle wind gust, caused by the predator's approach, excites cereal wind receptors, which encode both the presence and the direction of the stimulus. These cells in turn excite a group of giant interneurons (GI's) whose axons convey the directional information to thoracic motor centers. A given wind direction is coded not by a single GI functioning as a labeled line, but rather by some relationship among the spike trains in an assembly of GI's. This paper analyzes the code in this assembly. It is shown that all three pairs of GI's with the largest axonal diameters respond differentially to wind from left front vs. right front (Figs. 3, 4; Table 2). Each GI encodes these angles by both the time of its first action potential, and the number of action potentials, relative to its contralateral homolog. It is shown that the behavioral discrimination cannot rely solely upon the leftright differences in the time of the first action potential. A model of the assembly code is developed that involves a comparison of the numbers of action potentials in the left vs. the right group of giant interneurons. The model is shown to account for a large number of pre-existing experimental data on direction discrimination. The model requires, however, the involvement of additional cells in the left and right groups, besides the specific GI's whose role had been tested in prior experiments. The model is then tested by further experiments designed to verify the involvement of these added cells. These experiments support the model.