THE RECEPTIVE FIELDS OF RETINAL GANGLION CELLS: THE OUTPUT OF THE EYE Left: Four recordings from a typical on-center retinal ganglion cell. Each record is a single sweep of the oscilloscope, whose duration is 2.5 seconds. For a sweep this slow, the rising and falling phases of the impulse coalesce so that each spike appears as a vertical line. To the left the stimuli are shown. In the resting state at the top, there is no stimulus: firing is slow and more or less random. The lower three records show responses to a small (optimum size) spot, a large spot covering the receptive-field center and surround, and a ring covering the surround only. Right: Responses of an off-center retinal ganglion cell to the same set of stimuli shown at the left. In studying the retina we are confronted with two main problems: First, how do the rods and cones translate the light they receive into electrical, and then chemical, signals? Second, how do the subsequent cells in the next two layers-- the bipolar, horizontal, amacrine, and ganglion cells-- interpret this information? Before discussing the physiology of the receptors and intermediate cells, I want to jump ahead to describe the output of the retina-- represented by the activity of the ganglion cells. The map of the receptive field of a cell is a powerful and convenient shorthand description of the cell's behavior, and thus of its output. Understanding it can help us to understand why the cells in the intermediate stages are wired up as they are, and will help explain the purpose of the direct and indirect paths. If we know what ganglion cells are telling the brain, we will have gone far toward understanding the entire retina.