Cerebral Cortex, Vol 7, 523-533, Copyright © 1997 by Oxford University Press
H Markram
Tufted layer 5 (TL5) pyramidal neurons are important projection neurons
from the cerebral cortex to subcortical areas. Recent and ongoing
experiments aimed at understanding the computational analysis performed by
a network of synaptically connected TL5 neurons are reviewed here. The
experiments employed dual and triple whole-cell patch clamp recordings from
visually identified and preselected neurons in brain slices of
somatosensory cortex of young (14- to 16-day-old) rats. These studies
suggest that a local network of TL5 neurons within a cortical module of
diameter 300 microns consists of a few hundred neurons that are extensively
inter-connected with reciprocal feedback from at least first-, second- and
third-order target neurons. A statistical analysis of synaptic innervation
suggests that this recurrent network is not randomly arranged and hence
each neuron could be functionally unique. Synaptic transmission between
these neurons is characterized by use- dependent synaptic depression which
confers novel properties to this recurrent network of neurons. First, a
range of rates of depression for different synaptic connections enable each
TL5 neuron to receive a unique mixture of information about the average
firing rates and the temporally correlated action potential (AP) activity
in the population of presynaptic TL5 neurons. Second, each AP generated by
any neuron in the network induces a change (defined as an iteration step)
in the functional coupling of the neurons in the network (defined as
network configuration). It is proposed that the network configuration is
iterated during a stimulus to achieve an optimally orchestrated network
response. Hebbian, anti-Hebbian and neuromodulatory-induced modifications
of neurotransmitter release probability change the rates of synaptic
depression and thereby alter the iteration step size. These data may be
important to understand the dynamics of electrical activity within the
network.
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A network of tufted layer 5 pyramidal neurons
Department of Neurobiology, Weizmann Institute for Science, Rehovot, Israel. bnmark@weizmann.weizmann.ac.il
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