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1 Graduate School of Engineering Science, Osaka University, Toyonaka, Osaka, Japan
2 Graduate School of Frontier Biosciences, Osaka University, Toyonaka, Osaka, Japan
* To whom correspondence should be addressed. E-mail: ohzawa{at}fbs.osaka-u.ac.jp.
The receptive fields of complex cells in the early visual cortex are economically modeled by combining outputs of a quadrature pair of linear filters. For actual complex cells, such a minimal model may be insufficient, since many more simple cells are thought to make up a complex cell receptive field. To examine the minimalist model physiologically, we analyzed spatial relationships between the internal structure (subunits) and the overall receptive fields of individual complex cells by a two-stimulus interaction technique. The receptive fields of complex cells are more circular and only slightly larger than their subunits in size. In addition, complex cell subunits occupy similar spatial extents to simple cell receptive fields. Therefore, in these respects, the minimalist schema is a fair approximation to actual complex cells. However, there are violations against the minimal model. Simple cell receptive fields have significantly fewer subregions than complex cell subunits. And simple cell receptive fields are elongated more horizontally than vertically in general. This bias is absent in complex cell subunits and receptive fields. Thus, simple cells cannot be equated to individual complex cell subunits, and spatial pooling of simple cells may occur anisotropically to comprise a complex cell subunit. Moreover, when linear filters for complex cell subunits are examined separately for bright and dark responses, there are significant imbalances and position-displacements between them. This suggests that actual complex cell receptive fields are constructed by a richer combination of linear filters than proposed by the minimalist model.
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