Frequency coding is considered one of the most common coding strategies employed by neural systems. This fact leads, in experiments as well as in theoretical studies, to construction of so-called transfer functions, where the output firing frequency is plotted against the input intensity. The term firing frequency can be understood differently in different contexts. Basically, it means that the number of spikes over an interval of preselected length is counted and then divided by the length of the interval, but due to the obvious limitations, the length of observation cannot be arbitrarily long. Then firing frequency is defined as reciprocal to the mean interspike interval. In parallel, an instantaneous firing frequency can be defined as reciprocal to the length of current interspike interval, and by taking a mean of these, the definition can be extended to introduce the mean instantaneous firing frequency. All of these definitions of firing frequency are compared in an effort to contribute to a better understanding of the input-output properties of a neuron.
We present for the first time an analytical approach for determining the time of firing of multicomponent nonlinear stochastic neuronal models. We apply the theory of first exit times for Markov processes to the Fitzhugh-Nagumo system with a constant mean gaussian white noise input, representing stochastic excitation and inhibition. Partial differential equations are obtained for the moments of the time to first spike. The observation that the recovery variable barely changes in the prespike trajectory leads to an accurate one-dimensional approximation. For the moments of the time to reach threshold, this leads to ordinary differential equations that may be easily solved. Several analytical approaches are explored that involve perturbation expansions for large and small values of the noise parameter. For ranges of the parameters appropriate for these asymptotic methods, the perturbation solutions are used to establish the validity of the one-dimensional approximation for both small and large values of the noise parameter. Additional verification is obtained with the excellent agreement between the mean and variance of the firing time found by numerical solution of the differential equations for the one-dimensional approximation and those obtained by simulation of the solutions of the model stochastic differential equations. Such agreement extends to intermediate values of the noise parameter. For the mean time to threshold, we find maxima at small noise values that constitute a form of stochastic resonance. We also investigate the dependence of the mean firing time on the initial values of the voltage and recovery variables when the input current has zero mean.