Fig 4. Tuning of the differentiator readout network to implement the operation of the differentiator readout scheme.

A perturbation in the firing rate of the BCN (Δr) causes a perturbation in the mean input to the RS readout neurons, Δμ_e, and a perturbation in the firing rate of the inhibitory readout population $I$. This change in firing rate causes a shift in the input from $I$ to $S^B$ ($\Delta {\mu }_I$). The strength of the connection from $I$ to $S^B$ ($J_{ei}^R$) is adjusted such that $\Delta {\mu }_e+\Delta {\mu }_I\approx 0$. This cancellation reaches $S^B$ with a time lag ΔT so that the readout network roughly implements the operation of the DR Eq (4).