A quantitative model reveals a frequency ordering of prediction and prediction-error signals in the human brain

The human brain is proposed to harbor a hierarchical predictive coding neuronal network underlying perception, cognition, and action. In support of this theory, feedforward signals for prediction error have been reported. However, the identification of feedback prediction signals has been elusive due to their causal entanglement with prediction-error signals. Here, we use a quantitative model to decompose these signals in electroencephalography during an auditory task, and identify their spatio-spectral-temporal signatures across two functional hierarchies. Two prediction signals are identified in the period prior to the sensory input: a low-level signal representing the tone-to-tone transition in the high beta frequency band, and a high-level signal for the multi-tone sequence structure in the low beta band. Subsequently, prediction-error signals dependent on the prior predictions are found in the gamma band. Our findings reveal a frequency ordering of prediction signals and their hierarchical interactions with prediction-error signals supporting predictive coding theory.