Uncertainty-modulated prediction errors in cortical microcircuits

Understanding the variability of the environment is essential to function in everyday life. The brain must hence take uncertainty into account when updating its internal model of the world. The basis for updating the model are prediction errors that arise from a difference between the current model and new sensory experiences. Although prediction error neurons have been identified in layer 2/3 of diverse brain areas, how uncertainty modulates these errors and hence learning is, however, unclear. Here, we use a normative approach to derive how uncertainty should modulate prediction errors and postulate that layer 2/3 neurons represent uncertainty-modulated prediction errors (UPE). We further hypothesise that the layer 2/3 circuit calculates the UPE through the subtractive and divisive inhibition by different inhibitory cell types. By implementing the calculation of UPEs in a microcircuit model, we show that different cell types can compute the means and variances of the stimulus distribution. With local activity-dependent plasticity rules, these computations can be learned context-dependently, and allow the prediction of upcoming stimuli and their distribution. Finally, the mechanism enables an organism to optimise its learning strategy via adaptive learning rates.