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Thus complex spikes cannot only act as a teaching signal for a P-cell, but through complex spike synchrony, a P-cell population may act as a surrogate teacher for the DCN neuron that produced the erroneous output. It appears that grouping of P-cells into small populations that share a preference for error satisfies a critical requirement of efficient learning providing error information to the output layer neuron (DCN) that was responsible for the error, as well as the hidden layer neurons (P-cells) that contributed to it. This population
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