Natural scenes contain a variety of visual cues that facilitate boundary perception (e.g., luminance, contrast, and texture). Here we explore whether single neurons in early visual cortex can process both contrast and texture cues. We recorded neural responses in cat A18 to both illusory contours formed by abutting gratings (ICs, texture-defined) and contrast-modulated gratings (CMs, contrast-defined). We found that if a neuron responded to one of the two stimuli, it also responded to the other. These neurons signaled similar contour orientation, spatial frequency, and movement direction of the two stimuli. A given neuron also exhibited similar selectivity for spatial frequency of the fine, stationary grating components (carriers) of the stimuli. These results suggest that the cue-invariance of early cortical neurons extends to different kinds of texture or contrast cues, and might arise from a common nonlinear mechanism.

The discontinuities of phase-shifted abutting line gratings give rise to perception of an “illusory contour” (IC) along the line terminations. Neuronal responses to such ICs have been interpreted as evidence for a specialized visual mechanism, since such responses cannot be predicted from conventional linear receptive fields. However, when the spatial scale of the component gratings (carriers) is large compared to the neuron's luminance passband, these IC responses might be evoked simply by the luminance edges at the line terminations. Thus by presenting abutting gratings at a series of carrier spatial scales to cat A18 neurons, we were able to distinguish genuine nonlinear responses from those due to luminance edges. Around half of the neurons (both simple and complex types) showed a bimodal response pattern to abutting gratings: one peak at a low carrier spatial frequency range that overlapped with the luminance passband, and a second distinct peak at much higher frequencies beyond the neuron's grating resolution. For those bimodally responding neurons, the low-frequency responses were sensitive to carrier phase, but the high-frequency responses were phase-invariant. Thus the responses at low carrier spatial frequencies could be understood via a linear model, while the higher frequency responses represented genuine nonlinear IC processing. IC responsive neurons also demonstrated somewhat lower spatial preference to the periodic contours (envelopes) compared to gratings, but the optimal orientation and motion direction for both were quite similar. The nonlinear responses to ICs could be explained by the same energy mechanism underlying responses to second-order stimuli such as contrast-modulated gratings. Similar neuronal preferences for ICs and for gratings may contribute to the form-cue invariant perception of moving contours.

Cortical neurons selective for the direction of motion often exhibit some limited response to motion in their nonpreferred directions. Here we examine the dependence of neuronal direction selectivity on stimulus contrast, both for first-order (luminance-modulated, sine-wave grating) and second-order (contrast-modulated envelope) stimuli. We measured responses from single neurons in area 18 of cat visual cortex to both kinds of moving stimuli over a wide range of contrasts (1.25–80%). Direction-selective contrast response functions (CRFs) were calculated as the preferred-minus-null difference in average firing frequency as a function of contrast. We also applied receiver operating characteristic analysis to our CRF data to obtain neurometric functions characterizing the potential ability of each neuron to discriminate motion direction at each contrast level tested. CRFs for sine-wave gratings were usually monotonic; however, a substantial minority of neurons (35%) exhibited nonmonotonic CRFs (such that the degree of direction selectivity decreased with increasing contrast). The underlying preferred and nonpreferred direction CRFs were diverse, often having different shapes in a given neuron. Neurometric functions for direction discrimination showed a similar degree of heterogeneity, including instances of nonmonotonicity. For contrast-modulated stimuli, however, CRFs for either carrier or envelope contrast were always monotonic. In a given neuron, neurometric thresholds were typically much higher for second- than for first-order stimuli. These results demonstrate that the degree of a cell's direction selectivity depends on the contrast at which it is measured, and therefore is not a characteristic parameter of a neuron. In general, contrast response functions for first-order stimuli were very heterogeneous in shape and sensitivity, while those for second-order stimuli showed less sensitivity and were quite stereotyped in shape.