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Received April 8, 2003
Revised April 28, 2003
Accepted after revision May 22, 2003
1 Karolinska Institute
2 Karolinska Institutet
* To whom correspondence should be addressed. E-mail: yuri.zilberter{at}neuro.ki.se.
Neocortex, local networks, synaptic connections. The extent to which neocortical pyramidal cells function as a local network is determined by the strength and probability of their connections. By mapping connections between pyramidal cells we show that in a local network of about 600 pyramidal cells located within a cylindrical volume of 200 µm x 200 µm, of neocortical layer 2/3, an individual pyramidal cell receives synaptic inputs from about 60 other pyramidal neurons, with the majority of EPSP amplitudes in the 0.2- 1.0 mV range. The probability of connection decreases from 0.18 to 0.026 with inter-cell distance (over the range 25 µm to 200 µm). Within the same volume, local interneuron-pyramidal cell (fast spiking nonaccomodating interneuron, FS) connections are about ten times more numerous, with the majority of connections being reciprocal. The probability of excitatory and inhibitory connections between pyramidal cells and FS interneurons decreases only slightly with distance, being in the range 0.5-0.75. Pyramidal cells in the local network receive strong synaptic input during stimulation of afferent fibers in layers 1 and 6. Minimal-like stimulation of layer 1 or layer 6 inputs simultaneously induce post-synaptic potentials in connected pyramidal cells as well as in pyramidal-FS cell pairs. These inputs can readily induce firing of pyramidal cells, although synaptically connected cells display different firing patterns. Unitary EPSPs in pyramidal-pyramidal cell pairs do not detectably alter cell firing. FS interneurons fire simultaneously with pyramidal cells. In pyramidal-FS cell pairs, both unitary EPSPs and IPSPs efficiently modulate cell firing patterns. We suggest that computation in the local network may proceed not only by direct pyramidal-pyramidal communication, but also via local interneurons. With such a high degree of connectivity with surrounding pyramidal cells local interneurons are ideally poised to both coordinate and expand the local pyramidal cell network via pyramidal-interneuron-pyramidal communication.
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