A Simplified Model of Communication Between Time Cells: Accounting for the Linearly Increasing Timing Imprecision.

Many organisms can time intervals flexibly on average with high accuracy but substantial variability between the trials. One of the core psychophysical features of interval timing functions relates to the signatures of this timing variability; for a given individual, the standard deviation of timed responses/time estimates is nearly proportional to their central tendency (scalar property). Many studies have aimed at elucidating the neural basis of interval timing based on the neurocomputational principles in a fashion that would explain the scalar property. Recent experimental evidence shows that there is indeed a specialized neural system for timekeeping. This system, referred to as the "time cells," is composed of a group of neurons that fire sequentially as a function of elapsed time. Importantly, the time interval between consecutively firing time cell ensembles has been shown to increase with more elapsed time. However, when the subjective time is calculated by adding the distributions of time intervals between these sequentially firing time cell ensembles, the standard deviation would be compressed by the square root function. In light of this information the question becomes, "How should the signaling between the sequentially firing time cell ensembles be for the resulting variability to increase linearly with time as required by the scalar property?" We developed a simplified model of time cells that offers a mechanism for the synaptic communication of the sequentially firing neurons to address this ubiquitous property of interval timing. The model is composed of a single layer of time cells formulated in the form of integrate-and-fire neurons with feed-forward excitatory connections. The resulting behavior is simple neural wave activity. When this model is simulated with noisy conductances, the standard deviation of the time cell spike times increases proportionally to the mean of the spike-times. We demonstrate that this statistical property of the model outcomes is robustly observed even when the values of the key model parameters are varied.