Cholinergic Modulation Preserves Spike Timing Under Physiologically Realistic Fluctuating Input

Recently, there has been a vigorous debate concerning the nature of neural coding (Rieke et al. 1996; Stevens and Zador 1995; Shadlen and Newsome 1994). The prevailing viewhas been that the mean firing rate conveys all information about the sensory stimulus in a spike train and the precise timing of the individual spikes is noise. This belief is, in part, based on a lack of correlation between the precise timing ofthe spikes and the sensory qualities of the stimulus under study, particularly, on a lack of spike timing repeatability when identical stimulation is delivered. This view has been challenged by a number of recent studies, in which highly repeatable temporal patterns of spikes can be observed both in vivo (Bair and Koch 1996; Abeles et al. 1993) and in vitro (Mainen and Sejnowski 1994). Furthermore, application ofinformation theory to the coding problem in the frog and house fly (Bialek et al. 1991; Bialek and Rieke 1992) suggested that additional information could be extracted from spike timing. In the absence of direct evidence for a timing code in the cerebral cortex, the role of spike timing in neural coding remains controversial.