A method is developed for the analysis of nonlinear biological systems based on an input temporal signal that consists of a sum of a large number of sinusoids. Nonlinear properties of the system are manifest by responses at harmonics and intermodulation frequencies of the input frequencies. The frequency kernels derived from these nonlinear responses are similar to the Fourier transforms of the Wiener kernels. Guidelines for the choice of useful input frequency sets, and examples satisfying these guidelines, are given. A practical algorithm for varying the relative phases of the input sinusoids to separate high-order interactions is presented. The utility of this technique is demonstrated with data obtained from a cat retinal ganglion cell of the Y type. For a high spatial frequency grafting, the entire response is contained in the even-order nonlinear components. Even at low contrast, fourth-order components are detectable. This suggests the presence of an essential nonlinearity in the functional pathway of the Y cell, with its singularity at zero contrast.
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