Correlation versus gradient type motion detectors: the pros and cons

Visual motion contains a wealth of information about self-motion as well as the three- dimensional structure of the environment. Therefore, it is of utmost importance for any organism with eyes. However, visual motion information is not explicitly represented at the photoreceptor level, but rather has to be computed by the nervous system from the changing retinal images as one of the first processing steps. Two prominent models have been proposed to account for this neural computation: the Reichardt detector and the gradient detector. While the Reichardt detector correlates the luminance levels derived from two adjacent image points, the gradient detector provides an estimate of the local retinal image velocity by dividing the spatial and the temporal luminance gradient. As a consequence of their different internal processing structure, both the models differ in a number of functional aspects such as their dependence on the spatial-pattern structure as well as their sensitivity to photon noise. These different properties lead to the proposal that an ideal motion detector should be of Reichardt type at low luminance levels, but of gradient type at high luminance levels. However, experiments on the fly visual systems provided unambiguous evidence in favour of the Reichardt detector under all luminance conditions. Does this mean that the fly nervous system uses suboptimal computations, or is there a functional aspect missing in the optimality criterion? In the following, I will argue in favour of the latter, showing that Reichardt detectors have an automatic gain control allowing them to dynamically adjust their input-output relationships to the statistical range of velocities presented, while gradient detectors do not have this property. As a consequence, Reichardt detectors, but not gradient detectors, always provide a maximum amount of information about stimulus velocity over a large range of velocities. This important property might explain why Reichardt type of computations have been demonstrated to underlie the extraction of motion information in the fly visual system under all luminance levels.