Modeling of Neuronal Activity in the Visual Cortex

From Blasdel (J. of Neuroscience 12, 1992), the optical imaging of the orientation hypercolumn structure in monkey visual cortex.

From McLaughlin et al (PNAS 97, 2000), the response of a model visual cortex to oriented drifting grating stimulus.



An effective kinetic representation of fluctuation-driven neuronal networks with application to simple and complex cells in visual cortex
D. Cai, L. Tao, M. Shelley, and D. McLaughlin,
in PNAS 101, 7757-7762 (2004).

Abstract:  A coarse-grained representation of neuronal network dynamics is developed in terms of kinetic equations, which are derived by a moment closure, directly from the original large-scale integrate-and-fire (I&F) network. This powerful kinetic theory captures the full dynamic range of neuronal networks, from the mean-driven limit (a limit such as the number of neurons N -> {infty} , in which the fluctuations vanish) to the fluctuation-dominated limit (such as in small N networks). Comparison with full numerical simulations of the original I&F network establishes that the reduced dynamics is very accurate and numerically efficient over all dynamic ranges. Both analytical insights and scale-up of numerical representation can be achieved by this kinetic approach. Here, the theory is illustrated by a study of the dynamical properties of networks of various architectures, including excitatory and inhibitory neurons of both simple and complex type, which exhibit rich dynamic phenomena, such as, transitions to bistability and hysteresis, even in the presence of large fluctuations. The implication for possible connections between the structure of the bifurcations and the behavior of complex cells is discussed. Finally, I&F networks and kinetic theory are used to discuss orientation selectivity of complex cells for "ring-model" architectures that characterize changes in the response of neurons located from near "orientation pinwheel centers" to far from them.

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An Egalitarian Network Model for the Emergence of Simple and Complex Cells in Visual Cortex
L. Tao, M. Shelley, D. McLaughlin, and R. Shapley,
in PNAS 101, 366-371 (2004).

Abstract: We explain how Simple and Complex cells arise in a large-scale neuronal network model of the primary visual cortex of the macaque.  Our model consists of approximately 4,000 integrate-and-fire, conductance-based point neurons, representing the cells in a small, 1 square millimeter patch of an input layer of the primary visual cortex.  In the model the local connections are isotropic and nonspecific, and convergent input from the lateral geniculate nucleus confers cortical cells with orientation and spatial phase preference.  The balance between lateral connections and LGN drive determines whether individual neurons in this recurrent circuit are Simple or Complex.   The model reproduces qualitatively the experimentally observed distributions of both extracellular and intracellular measures of  Simple and Complex response.


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Mexican hats and pinwheels in Visual Cortex
K. Kang, M. Shelley, and H. Sompolinsky,
in PNAS 100, 2848-2853 (2003).

Abstract: Many models of cortical function assume that local lateral connections are specific with respect to the preferred features of the interacting cells and that they are organized in a Mexican-hat pattern with strong   center   excitation flanked by strong   surround   inhibition. However, anatomical data on primary visual cortex indicate that the local connections are isotropic and that inhibition has a shorter range than excitation. We address this issue in an analytical study of a neuronal network model of the local cortical circuit in primary visual cortex...

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States of High Conductance in a Large-Scale Model of the Visual Cortex

M. Shelley, D. McLaughlin, R. Shapley, and D.J. Wielaard

Journal of Computational Neuroscience 13,  pp. 93-109 (2002).

Abstract: This paper reports on the consequences of high, activity dependent, synaptic conductances in neurons in a large-scale neuronal network model of an input layer of the Macaque primary visual cortex (Area V1)....

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Coarse-Grained Reduction and Analysis of a Network Model of Cortical Response. I. Drifting Grating Stimuli

M. Shelley and D. McLaughlin

Journal of Computational Neuroscience 12, pp. 97-122 (2002)

Abstract:  We present a reduction of a large-scale network model of visual cortex developed by McLaughlin, Shapley, Shelley, and Wielaard.  The reduction is from many integrate-and-fire neurons to a spatially coarse-grained system for firing rates of neuronal subpopulations.  It accounts explicitly for ``disordered'' properties that vary widely from cortical neuron to cortical neuron, such as preferred spatial phase.  The result is a set of nonlinear spatio-temporal integral equations for ``phase-averaged'' firing rates across the model cortex.  For drifting grating stimuli, this system yields time invariant cortico-cortical conductances, with firing rates averaged over stimulus period being the natural objects.  Mathematical analysis then unveils the mechanisms underlying the spatially varying firing rates and orientation selectivity observed in the large-scale point-neuron simulations.  This reduction also reproduces, at far less computational cost, the salient features of the point-neuron network, and is used to study cortical response to changing stimulus contrast, noise level, and coupling length-scales.

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Efficient and Accurate Time-Integration Schemes for Integrate-and-Fire Neuronal Networks

M. Shelley and L. Tao

Journal of Computational Neuroscience 11, 111-119 (2001).

Abstract:  To avoid the numerical errors associated with resetting the potential following a spike in simulations of integrate-and-fire neuronal networks, Hansel et al. and Shelley independently developed a modified time-stepping method.  Their particular scheme consists of second-order Runge-Kutta time-stepping, a linear interpolant to find spike times, and a recalibration of post-spike potential using the spike times. Here we show analytically that such a scheme is second order, discuss the conditions under which efficient, higher-order algorithms can be constructed to treat resets, and develop a modified fourth-order scheme. To support our analysis, we simulate a system of integrate-and-fire conductance-based point neurons with all-to-all coupling.  For 6-digit accuracy, our modified Runge-Kutta fourth-order scheme needs a time-step of 10^{-3} seconds, whereas to achieve comparable accuracy using a recalibrated second-order or a first-order algorithm requires time-steps of 10^{-5} seconds or 10^{-9} seconds, respectively. Furthermore, since the cortico-cortical conductances in standard integrate-and-fire neuronal networks do not depend on the value of the membrane potential, we can attain fourth-order accuracy with computational costs normally associated with second-order schemes.

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How Simple Cells are made in a Nonlinear Network Model of the Visual Cortex

D.J. Wielaard, M.J. Shelley, David McLaughlin, and Robert Shapley

Journal of Neuroscience 21, pp. 5203-5211 (2001).

Abstract:

Simple cells in the striate cortex respond to visual stimuli in an approximately linear manner, even though the LGN input to the striate cortex, and the cortical network itself, are highly nonlinear...

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A Neuronal Network Model of Macaque Primary Visual Cortex (V1): 
Orientation Tuning and Dynamics in the Input Layer 4Calpha.

David McLaughlin, Robert Shapley, Michael Shelley, and D.J. Wielaard

PNAS (2000), v. 97, pp. 8087-8092

Abstract: Here, we offer an explanation for how selectivity for orientation could be produced by a model with circuitry that is based on the anatomy of V1 cortex....

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Computational Modeling of Orientation Tuning Dynamics in Monkey Primary Visual Cortex

M. C. Pugh, D. L. Ringach, R. Shapley and M. J. Shelley

Journal of Computational Neuroscience (2000), v. 8, pp. 143-159

Abstract: In the primate visual pathway, orientation tuning of neurons is first observed in the primary visual cortex. The LGN cells that comprise the thalamic input to V1 are not orientation tuned, but some V1 neurons are quite selective. Two main classes of theoretical models have been offered to explain orientation selectivity: feedforward models, in which inputs from spatially aligned LGN cells are summed together by one cortical neuron; and feedback models, in which an initial weak orientation bias due to convergent LGN input is sharpened and amplified by intracortical feedback. Recent data on the dynamics of orientation tuning, obtained by a cross-correlation technique, may help to distinguish between these classes of models. To test this possibility, we simulated the measurement of orientation tuning dynamics on various receptive field models, including a simple Hubel-Wiesel type feedforward model: a linear spatio-temporal filter followed by an integrate-and-fire spike generator. The computational study reveals that simple feedforward models may account for some aspects of the experimental data, but fail to explain many salient features of orientation tuning dynamics in V1 cells. A simple feedback model of interacting cells is also considered. This model is successful in explaining the appearance of Mexican-hat orientation profiles, but other features of the data continue to be unexplained.
 

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