Sensory areas should be adapted to the properties of their natural stimuli. What are the underlying rules that match the properties of complex cells in primary visual cortex to their natural stimuli? To address this issue, we sampled movies from a camera carried by a freely moving cat, capturing the dynamics of image motion as the animal explores an outdoor environment. We use these movie sequences as input to simulated neurons. Following the intuition that many meaningful high-level variables, e.g., identities of visible objects, do not change rapidly in natural visual stimuli, we adapt the neurons to exhibit firing rates that are stable over time. We find that simulated neurons, which have optimally stable activity, display many properties that are observed for cortical complex cells. Their response is invariant with respect to stimulus translation and reversal of contrast polarity. Furthermore, spatial frequency selectivity and the aspect ratio of the receptive field quantitatively match the experimentally observed characteristics of complex cells. Hence, the population of complex cells in the primary visual cortex can be described as forming an optimally stable representation of natural stimuli.
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