A, linear model of simple cells. The first stage is linear filtering, i. e. aweighted sum of the image intensities, with weights given by the receptive field. The second stage isrectification: only the part of the responses that is larger than a threshold is seen in the firing rate response. B, subunit model of complex cells. The first stage is linear filtering by a number of receptive fields such asthose of simple cells (here we show four of them with spatial phases offset by 90 deg). The subsequent stages involve rectification, and then summation.