Analysis of multidimensional difference-of-Gaussians filters in terms of directly observable parameters.

The difference-of-Gaussians (DOG) filter is a widely used model for the receptive field of neurons in the retina and lateral geniculate nucleus (LGN) and is a potential model in general for responses modulated by an excitatory center with an inhibitory surrounding region. A DOG filter is defined by three standard parameters: the center and surround sigmas (which define the variance of the radially symmetric Gaussians) and the balance (which defines the linear combination of the two Gaussians). These parameters are not directly observable and are typically determined by nonlinear parameter estimation methods applied to the frequency response function. DOG filters show both low-pass (optimal response at zero frequency) and bandpass (optimal response at a nonzero frequency) behavior. This paper reformulates the DOG filter in terms of a directly observable parameter, the zero-crossing radius, and two new (but not directly observable) parameters. In the two-dimensional parameter space, the exact region corresponding to bandpass behavior is determined. A detailed description of the frequency response characteristics of the DOG filter is obtained. It is also found that the directly observable optimal frequency and optimal gain (the ratio of the response at optimal frequency to the response at zero frequency) provide an alternate coordinate system for the bandpass region. Altogether, the DOG filter and its three standard implicit parameters can be determined by three directly observable values. The two-dimensional bandpass region is a potential tool for the analysis of populations of DOG filters (for example, populations of neurons in the retina or LGN), because the clustering of points in this parameter space may indicate an underlying organizational principle. This paper concentrates on circular Gaussians, but the results generalize to multidimensional radially symmetric Gaussians and are given as an appendix.