An examination of physiological mechanisms underlying the frequency-doubling illusion.

PURPOSE The frequency-doubling illusion is an apparent doubling of spatial frequency when a sinusoidal grating is modulated rapidly in temporal counterphase. It has been proposed that the illusion arises from a spatially nonlinear ganglion cell class. The current study reexamines this possibility and investigates other mechanisms that may underlie the illusion. METHODS Responses of macaque magnocellular (MC) retinal ganglion cells were recorded to counterphase-modulated sinusoidal gratings of various spatial frequencies, and linearity of spatial summation was assessed. Human psychophysical thresholds were measured for a variety of phase discrimination and matching tasks. RESULTS Consistent with lateral geniculate recordings reported by other authors, no evidence was found of a separate nonlinear (M(y)) MC cell class. The small, spatially nonlinear responses found were least at the low spatial frequencies used in clinical testing. Further analysis showed that no spatially modulated signal can be expected from the nonlinear response of a ganglion cell; the nonlinearity of spatial summation gives a doubled response in time but not across space. Psychophysical performance was consistent with an inability to distinguish the temporal phase of counterphase-modulated gratings when the illusion occurs. From 4 to 40 Hz, the zero-crossings of the modulated sinusoidal grating provided a spatial cue and were matched to comparison patterns at twice the stimulus spatial frequency. CONCLUSIONS These results are inconsistent with the hypothesis that spatially nonlinear (M(y)) retinal ganglion cells are the physiological substrate of the frequency-doubling illusion. A cortical loss of temporal phase discrimination may be the principle cause of the illusion.