Analysis of integrate-and-fire neurons: synchronization of synaptic input and spike output.

A new technique for analyzing the probability distribution of output spikes for the integrate-and-fire model is presented. This technique enables us to investigate models with arbitrary synaptic response functions that incorporate both leakage across the membrane and a rise time of the postsynaptic potential. The results, which are compared with numerical simulations, are exact in the limit of a large number of small-amplitude inputs. This method is applied to the synchronization problem, in which we examine the relationship between the spread in arrival times of the inputs (the temporal jitter of the synaptic input) and the resultant spread in the times at which the output spikes are generated (output jitter). The results of previous studies, which indicated that the ration of the output jitter to the input jitter is consistently less than one and that it decreases for increasing numbers of inputs, are confirmed for three classes of the integrate-and-fire model. In addition to the previously identified factors of axonal propagation times and synaptic jitter, we identify the variation in the spike-generating thresholds of the neurons and the variation in the number of active inputs as being important factors that determine the timing jitter in layered networks. Previously observed phase differences between optimally and suboptimally stimulated neurons may be understood in terms of the relative time taken to reach threshold.