Visual cortex combines a stimulus and an error-like signal with a proportion that is dependent on time, space, and stimulus contrast

David Eriksson, Thomas Wunderle, Kerstin Schmidt

Even though the visual cortex is one of the most studied brain areas, the neuronal code in this area is still not fully understood. In the literature, two codes are commonly hypothesized, namely stimulus and predictive (error) codes. Here, we examined whether and how these two codes can coexist in a neuron. To this end, we assumed that neurons could predict a constant stimulus across time or space, since this is the most fundamental type of prediction. Prediction was examined in time using electrophysiology and voltage-sensitive dye imaging in the supragranular layers in area 18 of the anesthetized cat, and in space using a computer model. The distinction into stimulus and error code was made by means of the orientation tuning of the recorded unit. The stimulus was constructed as such that a maximum response to the non-preferred orientation indicated an error signal, and the maximum response to the preferred orientation indicated a stimulus signal. We demonstrate that a single neuron combines stimulus and error-like coding. In addition, we observed that the duration of the error coding varies as a function of stimulus contrast. For low contrast the error-like coding was prolonged by around 60-100%. Finally, the combination of stimulus and error leads to a suboptimal free energy in a recent predictive coding model. We therefore suggest a straightforward modification that can be applied to the free energy model and other predictive coding models. Combining stimulus and error might be advantageous because the stimulus code enables a direct stimulus recognition that is free of assumptions whereas the error code enables an experience dependent inference of ambiguous and non-salient stimuli.