Two-layer model with feedback.

<p>(A) Feedback modulates the subunit inputs by an additive or multiplicative term (indicated by the placeholder #). These feedback terms are non-linear functions (<i>g</i><sub>1</sub> and <i>g</i><sub>2</sub>) of the response. (B-D) Impact of feedback on the response for the two-layer model shown in <a href="http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1006757#pcbi.1006757.g003" target="_blank">Fig 3D</a>. (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 (<a href="http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1006757#pcbi.1006757.g003" target="_blank">Fig 3D</a>). 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.</p>