We present a neural model for how the Mauthner system could compute the direction of a transient sound stimulus originating on either the left or right side of a fish. This computation results in an initial orientation of an escape response away from the side of the stimulus. Our idea is based on the phase model of underwater sound localization by fishes. If the phase model is applicable to the Mauthner system, then the problem of sound localization can be reduced to a logical operator, the EXCLUSIVE-NOR (or XNOR). We show how this can be solved by the Mauthner system using afferents that convey separate inputs of sound pressure transduced by the swimbladder (rarefaction and compression) and particle displacement (left and right) from the inner ear. In our model, both pressure components are responsible for bringing the Mauthner cell to threshold. Mauthner firing is gated by the inhibitory PHP neurons receiving specific combinations of pressure and displacement that implement the XNOR logic. We refer to this as the XNOR model. This model is experimentally verifiable and makes specific predictions about the expected acoustic response characteristics of the Mauthner and PHP neurons. Our model places a component of PHP function into a new neuroethological context and may provide insights into the central neurophysiological mechanisms of directional hearing in fishes. In particular, we show how the XNOR model can be applied to predict the activity of diverse neural elements involved in acoustic localization by fishes.

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