Shunting inhibition, a silent step in visual cortical computation

Brain computation, in the early visual system, is often considered as a hierarchical process in which features extracted in a given sensory relay are not present in previous stages of integration. In particular, orientation preference and its fine tuning selectivity are functional properties shared by most cortical cells and they are not observed at the preceding geniculate stage. A classical problem is identifying the mechanisms and circuitry underlying these computations. Several organizational principles have been proposed, giving different weights to the feedforward thalamocortical drive or to intracortical recurrent architectures. Within this context, an important issue is whether intracortical inhibition is fundamental for the genesis of stimulus selectivity, or rather normalizes spike response tuning with respect to other features such as stimulus strength or contrast, without influencing the selectivity bias and preference expressed in the excitatory input alone. We review here experimental observations concerning the presence or absence of inhibitory input evoked by non-preferred orientation/directions. Intracellular current clamp and voltage clamp recordings are analyzed in the light of new methods allowing us (1) to increase the visibility of inhibitory input, and (2) to continuously measure the visually evoked dynamics of input conductances. We conclude that there exists a diversity of synaptic input combinations generating the same profile of spike-based orientation selectivity, and that this diversity most likely reflects anatomical non-homogeneities in input sampling provided by the local context of the columnar and lateral intracortical network in which the considered cortical cell is embedded.