Of all sensory cortical areas, barrel cortex is among the best understood in terms of circuitry, yet least understood in terms of sensory function. Because sensory cortical areas have stereotyped anatomies, understanding computations in one sensory area may inform us of computations being performed by other sensory areas or sensory microcircuits all over the brain. Functional studies of barrel cortex are therefore important for marrying our immense and increasing knowledge of the cortical circuitry with the computations being performed in a cortical microcircuit. This thesis is an investigation of barrel cortex function as it pertains to 1) site specific sensory evoked plasticity in cortical microcircuit and 2) sensory receptive fields of the different cortical lamina in S1. The brain's capacity to rewire is thought to diminish with age. It is widely believed that development stabilizes the synapses from thalamus to cortex and that adult experience alters only synaptic connections between cortical neurons.

We combined whole-cell recording from individual thalamocortical neurons in adult rats with a newly developed automatic tracing technique to reconstruct individual axonal trees. Whisker trimming substantially reduced thalamocortical axon length in barrel cortex but not the density of TC synapses along a fiber. Thus, sensory experience alters the total number of TC synapses. After trimming, sensory stimulation evoked more tightly time-locked responses among thalamorecipient layer 4 cortical neurons. Axonal plasticity was topographically specific, with robust changes in L4 and modest changes in the septal and infragranular layers. These results indicate that plasticity is mediated by interactions with the local cortical subcircuit and may be suggestive of laminar specific roles in sensory learning/coding. Next we sought to examine spatiotemporal coding properties of neurons in the different layers of the cortical microcircuit in S1. We combined intracellular recording and a novel multi-directional multi-whisker stimulator system to estimate receptive fields by reverse correlation of stimuli to synaptic inputs.

Spatiotemporal receptive fields were identified orders of magnitude faster than by conventional spike-based approaches, even for neurons with little or no spiking activity. Given a suitable stimulus representation, a simple linear model captured the stimulus-response relationship for all neurons with unprecedented accuracy. In contrast to conventional single-whisker stimuli, complex stimuli revealed dramatically sharpened receptive fields, largely due to the effects of adaptation. Surprisingly, this phenomenon allows the surround to facilitate rather than suppress responses to the principal whisker. Optimized stimuli enhanced firing in layers 4-6, but not 2/3, which remained sparsely active. Surround facilitation through adaptation may be required for discriminating complex shapes and textures during natural sensing.