Intracellular measurements of spatial integration and the MAX operation in complex cells of the cat primary visual cortex

We have examined the spatial integration properties of complex cells to determine whether some of their responses can be described by a maximum operation (MAX)-like computation, as suggested by Riesenhuber and Poggio's model of object recognition. Membrane potential was recorded from anesthetized cats while optimally oriented bars were presented, either alone or in pairs, in different parts of the cells' receptive field. In most cells, the membrane potential response to two bars presented simultaneously could not be predicted by the sum of the responses to individual bars. In many cells, however, the responses closely approximated a MAX-like model. That is, the response of the cell to two bars was similar to the larger of the two individual responses (soft-MAX). The degree of nonlinear summation varied from cell to cell and varied within single cells from one stimulus configuration to another but on average fit most closely to the MAX model. The firing response of the cells was also well predicted by the MAX-like model. The MAX-like behavior was independent of the distance between the bars (orthogonal to the preferred orientation), independent of the relative amplitude of the responses, and slightly less pronounced at low levels of contrast. This MAX-like behavior of a subset of complex cells may play an important role in invariant object recognition in clutter.