Two-layer model with feedback.

(A) Feedback modulates the subunit inputs by an additive or multiplicative term (indicated by the placeholder #). These feedback terms are non-linear functions (g1 and g2) of the response. (B-D) Impact of feedback on the response for the two-layer model shown in Fig 3D. (B) Negative feedback shifts the iso-response curves further apart. (C) Positive feedback shifts the iso-response curves closer together. (D) Multiplicative feedback rescales the iso-response curves. Top insets show iso-response curves from above. Dashed lines illustrate the purple iso-response curve of the two-layer model without feedback (Fig 3D). Bottom and right insets show subunit non-linearities and feedback functions. The subunit non-linearities of the two-layer models without feedback are illustrated by the blue and the dark red dashed lines. Solid lines indicate the subunit non-linearities of the corresponding two-layer models with feedback for stimuli on three iso-response curves (black, purple and yellow lines). The feedback is constant on iso-response curves and shifts and rescales the subunit non-linearities. On the gray iso-response curves the feedback is zero.