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The Journal of Neuroscience, July 15, 2001, 21(14):5203-5211
How Simple Cells Are Made in a Nonlinear Network Model of the
Visual Cortex
D. J.
Wielaard,
Michael
Shelley,
David
McLaughlin, and
Robert
Shapley
Center for Neural Science and Courant Institute of Mathematical
Sciences, New York University, New York, New York 10012
Simple cells in the striate cortex respond to visual stimuli in an
approximately linear manner, although the LGN input to the striate
cortex, and the cortical network itself, are highly nonlinear. Although
simple cells are vital for visual perception, there has been no
satisfactory explanation of how they are produced in the cortex. To
examine this question, we have developed a large-scale neuronal network
model of layer 4C in V1 of the macaque cortex that is
based on, and constrained by, realistic cortical anatomy and
physiology. This paper has two aims: (1) to show that neurons in the
model respond like simple cells. (2) To identify how the model
generates this linearized response in a nonlinear network. Each neuron
in the model receives nonlinear excitation from the lateral geniculate
nucleus (LGN). The cells of the model receive strong (nonlinear)
lateral inhibition from other neurons in the model cortex. Mathematical
analysis of the dependence of membrane potential on synaptic
conductances, and computer simulations, reveal that the nonlinearity of
corticocortical inhibition cancels the nonlinear excitatory input from
the LGN. This interaction produces linearized responses that agree with
both extracellular and intracellular measurements. The model correctly
accounts for experimental results about the time course of simple cell
responses and also generates testable predictions about variation in
linearity with position in the cortex, and the effect on the linearity
of signal summation, caused by unbalancing the relative strengths of
excitation and inhibition pharmacologically or with extrinsic current.
Key words:
primary visual cortex; neuronal network model; simple
cells; linearity; synaptic inhibition; phase averaging
Copyright © 2001 Society for Neuroscience 0270-6474/01/21145203-09$05.00/0
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