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Direction selectivity (DS) of neuronal responses is fundamental for motion detection. How the integration of synaptic excitation and inhibition contributes to DS however remains not well-understood. Here, in vivo whole-cell voltage-clamp recordings in mouse primary visual cortex (V1) revealed that layer 4 simple cells received direction-tuned excitatory inputs but barely tuned inhibitory inputs under drifting-bar stimulation. Excitation and inhibition exhibited differential temporal offsets under movements of opposite directions: excitation peaked earlier than inhibition at the preferred direction, and vice versa at the null direction. This could be attributed to a small spatial mismatch between overlapping excitatory and inhibitory receptive fields: the distribution of excitatory input strengths was skewed and the skewness was strongly correlated with the DS of excitatory input, whereas that of inhibitory input strengths was spatially symmetric. Neural modeling revealed that the relatively stronger inhibition under null directional movements, as well as the specific spatial-temporal offsets between excitation and inhibition, allowed inhibition to enhance the DS of output responses by suppressing the null response more effectively than the preferred response. Our data demonstrate that while tuned excitatory input provides the basis for DS in mouse V1, the largely untuned and spatiotemporally offset inhibition contributes importantly to sharpening of DS.

Strengthening of Direction Selectivity by Broadly Tuned and Spatiotemporally Slightly Offset Inhibition in Mouse Visual Cortex