Processing of second-order stimuli in the visual cortex

Naturally occurring visual stimuli are rich in examples of objects delineated from their backgrounds simply by differences in luminance, so-called first-order stimuli, as well as those defined by differences of contrast or texture, referred to as second-order stimuli. Here we provide a brief overview of visual cortical processing of second-order stimuli, as well as some comparative background on first-order processing, concentrating on single-unit neurophysiology, but also discussing relationships to human psychophysics and to neuroimaging. The selectivity of visual cortical neurons to orientation, spatial frequency, and direction of movement of first-order, luminance-defined stimuli is conventionally understood in terms of simple linear filter models, albeit with some minor nonlinearities such as thresholding and gain control. However, these kinds of models fail entirely to account for responses of neurons to second-order stimuli such as contrast envelopes, illusory contours, or texture borders. Second-order stimuli constructed from sinusoidal components have been used to analyze the neurophysiological mechanisms of such responses; these experiments demonstrate that the same neuron can exhibit three distinct kinds of tuning to spatial frequency, and also to orientation. These results can be understood in terms of a type of nonlinear 'filter-->rectify-->filter' model, which has been widely used in human psychophysics. Finally, several general issues will be discussed, including potential artifacts in experiments with second-order stimuli, and strategies for avoiding or controlling for them; caveats about definitions of first- vs. second-order mechanisms and stimuli; the concept of form-cue invariance; and the functional significance of second-order processing.