Modeling corticofugal feedback and the sensitivity of lateral geniculate neurons to orientation discontinuity

We model feedback from primary visual cortex to the dorsal lateral geniculate nucleus (dLGN). This feedback makes dLGN neurons sensitive to orientation discontinuity (Sillito et al., 1993; Cudeiro & Sillito, 1996). In the model, each dLGN neuron receives retinotopic input driven by layer 6 cortical neurons in a full set of orientation columns. Excitation is monosynaptic, while inhibition is through perigeniculate neurons and dLGN interneurons. The stimulus consists of drifting gratings, one within and the other outside a circular region centered over the receptive field of the model dLGN relay neuron we study. They appear as a single grating when they are aligned with equal contrast. The model reproduces experimental results showing an increasing inhibitory effect of feedback on the firing rate of dLGN neurons as the two gratings move towards the aligned position. Moreover, enhancement of dLGN cell center-surround antagonism by feedback is revealed by measuring the responses to drifting gratings inside a circular window, as a function of window radius. This effect is related to the observed length tuning of dLGN cells. Sensitivity to orientation discontinuity could be mediated in the model by feedback from either simple or complex cells. The model puts constraints on the feedback synaptic footprint and shows that its elongated shape does not play a crucial role in sensitivity to orientation discontinuity. The inhibitory component of feedback must predominate overall, but the feedback signal from a cortical neuron to a dLGN neuron with the same or nearby receptive-field center can be dominated by excitation. Predictions of the model include (1) robust stimuli for layer 6 cortical neurons give pronounced nonlinearities in the responses of dLGN neurons; (2) the sensitivity to orientation discontinuity at low contrast is twice that at high contrast.