Recently proposed models of orientation tuning in layer 4 of cat primary visual cortex (Somers, Nelson, & Sur, 1995; Douglas, Koch, Mahowald, Martin, & Suarez, 1995) rely on widespread inhibitory intracortical connections to suppress the nonoptimal component of a broadly tuned thalamic input, while local excitatory intracortical connections amplify the optimal component. However, new experimental data (Ferster, Chung, & Wheat, 1996) and theoretical analyses (Ferster, 1987; Krukowski, Priebe, & Miller, 1996) show that the temporally modulated component of thalamic input is well tuned and that the cortical circuitry must simply subtract an unmodulated DC component at nonoptimal orientations to obtain sharp tuning. In addition, within a single hypercolumn in layer 4, inhibitory and excitatory layer 4 neurons have approximately equal-sized axonal fields, making the most of their synapses within their own dendritic field (Kisvarday, Martin, Whitteridge, & Somogyi, 1985; Martin & Whitteridge, 1984). We have constructed a model of a single microcolumn in which GABAB inhibition subtracts the DC and controls the sustained response, while GABAA inhibition controls the response to transient and suprathreshold inputs. The model fits experimental data based on stimulation with drifting sine-wave gratings as well as flashed bars, explains a counterintuitive property of the GABAB K+ conductance, and at suboptimal orientations and submaximal contrasts produces an exponential distribution of firing frequencies.

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