Whisker sensory system - from receptor to decision.

One of the great challenges of systems neuroscience is to understand how the neocortex transforms neuronal representations of the physical characteristics of sensory stimuli into the percepts which can guide the animal's decisions. Here we present progress made in understanding behavioral and neurophysiological aspects of a highly efficient sensory apparatus, the rat whisker system. Beginning with the 1970s discovery of "barrels" in the rat and mouse brain, one line of research has focused on unraveling the circuits that transmit information from the whiskers to the sensory cortex, together with the cellular mechanisms that underlie sensory responses. A second, more recent line of research has focused on tactile psychophysics, that is, quantification of the behavioral capacities supported by whisker sensation. The opportunity to join these two lines of investigation makes whisker-mediated sensation an exciting platform for the study of the neuronal bases of perception and decision-making. Even more appealing is the beginning-to-end prospective offered by this system: the inquiry can start at the level of the sensory receptor and conclude with the animal's choice. We argue that rats can switch between two modes of operation of the whisker sensory system: (1) generative mode and (2) receptive mode. In the generative mode, the rat moves its whiskers forward and backward to actively seek contact with objects and to palpate the object after initial contact. In the receptive mode, the rat immobilizes its whiskers to optimize the collection of signals from an object that is moving by its own power. We describe behavioral tasks that rats perform in these different modes. Next, we explore which neuronal codes in sensory cortex account for the rats' discrimination capacities. Finally, we present hypotheses for mechanisms through which "downstream" brain regions may read out the activity of sensory cortex in order to extract the significance of sensory stimuli and, ultimately, to select the appropriate action.