We have used frequency-domain methods to characterize the spatial receptive-field structure of cat retinal W cells. For most ON- and OFF-center tonic and phasic W cells, measurements of responsivity to drifting gratings at various spatial frequencies could be adequately described by a difference-of-Gaussians (DOG) function, consistent with the presence of center and surround mechanisms that are approximately Gaussian in shape and whose signals are combined additively. Estimates of the responsivity of the center mechanisms of tonic and phasic W cells were similar, but both were significantly lower than the corresponding values for X or Y cells. The width of the center mechanisms of tonic W cells, phasic W cells, and Y cells did not differ significantly from each other, but all were significantly larger than the width of X-cell centers. Surround parameters did not vary significantly among the four groups of ganglion cells. Measurements of contrast gain in both tonic and phasic W cells gave values that were significantly lower than in X or Y cells.
Virtually all of the phasic W cells in our sample displayed evidence of spatial non-linearities in their receptive fields, in the form of either d.c. responses to drifting sine-wave gratings or second harmonic responses to counterphased gratings. The spatial resolution of the mechanism underlying these nonlinearities was typically higher than that of the center mechanism of these cells. Most tonic W cells exhibited linear spatial summation, although a subset gave strong second harmonic responses to counterphased gratings.
Spatial-responsivity measurements for most ON-OFF and directionally selective W cells were not adequately described by DOG functions. These cells did, however, show evidence of spatial nonlinearities similar to those seen in phasic W cells. Suppressed-by-contrast cells gave both modulated and unmodulated responses to drifting gratings which both appeared to involved rectification, but which differed from each other in both spatial resolution and contrast gain.
These data confirm earlier reports that the receptive fields of tonic and most ON- or OFF-center phasic W cells appear to include classical center and surround mechanisms. However, the receptive fields of some phasic cells, as well as ON-OFF and directionally selective W cells may have quite different structures. Our results also suggest that phasic, ON-OFF, directionally selective, suppressed-by-contrast, and a subset of tonic W cells may all receive nonlinear inputs with characteristics similar to those described in the receptive fields of retinal Y cells. If so, this has important implications for identifying and understanding the presynaptic circuitry of W cells, as well as the nature of their output to both telencephalic and midbrain visual targets.