Numerous careful behavioral studies of visual persistence have reported a variety of apparently contradictory effects. Variation of flash intensity has particularly been found to have both direct and inverse effects on subjective duration. This conflict has been addressed by theories which contain both sensory and cognitive components. Depending on the weight given to these components, one obtains theories which emphasize intensity dependence or task dependence. Few comparably detailed physiological studies of persistence exist. To clarify the issues raised by these theories, we examined the responses evoked in the model photoreceptor of the Limulus lateral eye. To explore the role of sensory variables, we manipulated adaptation state and flash intensity. To explore cognitive variables, the durations of the photoreceptor potentials (RPs) evoked in this model system were assessed by a mutually complementary and complete set of candidate sensory codes. Accordingly, sharp microelectrodes were used to record RPs intracellularly from single photoreceptor cells in response to 40-ms flashes whose intensity was varied over at least 3.6 log units. Two light adaptation states were used which differed in sensitivity by 3.5 log units. This model system made it possible to employ stringent objective assessments of data quality which ensured that only cells which had remained stable for several hours contributed to the present data. A variety of code-dependent trends were found: direct, inverse, invariant, and U-shaped trends related flash intensity to RP duration, while adaptation state interacted with some of these trends. Only some of the expectations which had generated this research were qualitatively corroborated and numerous quantitative discrepancies were found between data and theory. While caution is necessary when generalizing from neural responses to perception, these data indicate that two major gaps now exist in this field. First, both task and stimulus variables need to be exhaustively explored in more complete behavioral experiments. The present data make it more likely that sensory models and cognitive models simply address different aspects of the same phenomenon. Second, similarly detailed questions need to be posed to more central neural loci, particularly to those in the various visual cortices.

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Perceived duration can be assessed behaviorally by adjusting the interval between two flashes so that an observer just perceives a certain relation between them. In such studies, the cognitive characteristics of the required relation necessarily interact with the sensory characteristics of the responses evoked by the two flashes. To dissociate the contributions of these two factors, we executed a physiological study which yielded more complete information on the role of each factor in two paradigms which have been used to characterize perceived duration behaviorally, namely the persistence-of-form design and the successive field design. The effect of sensory manipulations have yielded particularly problematic results in these two paradigms because opposite trends were found when intensity was varied. Intracellular recordings were therefore taken from photoreceptor cells exposed to procedural manipulations which match the sensory and cognitive variations employed in behavioral paradigms. The sensory variables of flash intensity, state of adaptation, and flash interval were explored with some completeness. Cognitive factors were assessed in two ways. First, the contribution of the neural site of sensory integration was determined by making a clear distinction between data collected when all stimuli affect the same receptors versus data collected when different stimuli affect different receptors. Second, the consequences of arbitrary choices of candidate code and dependent variable were also explored. When so organized, the physiological data provide a coherent basis for harmonizing apparently contradictory behavioral results because they qualitatively paralleled the behavioral data's complex dependence on intensity and interval. In particular, both direct and inverse dependencies of response duration on intensity exist in both physiology and behavior with the exact nature of the trend depending as much on the cognitive analysis of the neural responses as on their dynamics and energetics. Further, large quantitative differences were found which also were an expression of the different ways in which the two behavioral paradigms affect receptor potentials.