The direction-selective contrast response of area 18 neurons is different for first- and second-order motion

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.