Areal influences on complex cells in cat striate cortex: stimulus-specificity of width and length summation

In single neurones recorded from the striate cortex of cats anaesthetized with N2O/O2/halothane, receptive field dimensions, length specificity and areal extent of drive were assessed for different classes of visual stimuli. Receptive fields were mapped as rectangular minimum response fields (MRFs). Spatial summation along the axis of preferred orientation was assessed: for moving bars whose length was varied (length summation); and for height variation of a square-wave grating patch against a uniform grey background, or a patch of moving texture against a stationary background of similar texture. In complementary tests a moving square-wave grating background was progressively occluded by a uniform grey foreground mask of variable height; or a mask of stationary texture of variable height progressively occluded a background of moving texture. In parallel measurements, the width of grating or textured patches or masks was varied whilst maintaining height constant. Broadly speaking, the areal influence of each class of stimulus was comparable, and distinct from extra-receptive field phenomena in evoking responses from within the receptive field, but not from surrounding areas. The masking paradigm provided the most sensitive measure of receptive field height and width. However, in some neurones length summation, the degree of end-stopping, and the directional bias depended critically on the stimulus configuration used. Length summation tended to be more dramatic for short bars than for gratings. Length summation for texture was significantly more pronounced than for an oriented bar in special and in intermediate complex neurones. By contrast, endstopping was typically less intense for gratings than for bars, and least pronounced for texture. Because of stimulus specificity, complex neurones assigned to particular functional subgroups on the basis of their response to oriented bars may exhibit quite different patterns of behaviour for other classes of stimuli.