Assessing the reliability of neuronal spike trains is fundamental to an understanding of the neural code. We measured the reproducibility of retinal responses to repeated visual stimuli. In both tiger salamander and rabbit, the retinal ganglion cells responded to random f licker with discrete, brief periods of firing. For any given cell, these firing events covered only a small fraction of the total stimulus time, often less than 5%. Firing events were very reproducible from trial to trial: the timing jitter of individual spikes was as low as 1 msec, and the standard deviation in spike count was often less than 0.5 spikes. Comparing the precision of spike timing to that of the spike count showed that the timing of a firing event conveyed several times more visual information than its spike count. This sparseness and precision were general characteristics of ganglion cell responses, maintained over the broad ensemble of stimulus waveforms produced by random f licker, and over a range of contrasts. Thus, the responses of retinal ganglion cells are not properly described by a firing probability that varies continuously with the stimulus. Instead, these neurons elicit discrete firing events that may be the fundamental coding symbols in retinal spike trains.All of our visual experience derives from sequences of action potentials traveling down the optic nerve. Many theories have been proposed to explain how these spike trains from retinal ganglion cells encode the visual world (1-4). Fundamental to such an understanding is the reproducibility of these neural symbols to repeated presentations of the same stimulus. If retinal spike trains are highly deterministic, then individual visual messages can be attached to each spike; whereas, if they are highly stochastic, then the brain must average over many spikes to obtain an equally informative visual message.As far back as 1928, Adrian (5) proposed that information about the sensory environment is conveyed in the time-varying firing rate of spiking sensory neurons-a view that has been influential to neuroscience ever since (6-8). As a result, many researchers have concentrated on estimates of the firing rate derived from averages over long time windows or multiple stimulus presentations (9, 10). Measurements of response reliability have often focused on the trial-to-trial variance in this spike count: in the visual cortex, this variance is found to be greater than the mean (11, 12), whereas similar experiments in the thalamus and retina have found variance-to-mean ratios both above and below one (13-15). The picture emerging from this work is that spike trains in the visual system are intrinsically stochastic; that, at best, one can determine the instantaneous probability that the neuron will fire, and that this firing rate depends in some smooth fashion on the sensory stimulus. However, poor reproducibility can also arise from confounding factors (16), such as anesthesia (13), uncontrolled eye movements (17, 18), or ongoing brain activity (19). Furthermore, the ...