Forward- and backpropagation in a silicon dendrite

We have developed an analog very-large-scale integrated (aVLSI) electronic circuit that emulates a compartmental model of a neuronal dendrite. The horizontal conductances of the compartmental model are implemented as a switched capacitor network. The transmembrane conductances are implemented as transconductance amplifiers. The electrotonic properties of our silicon cable are qualitatively similar to those of the ideal passive cable that is commonly used to model mathematically the electrotonic behavior of neurons. In particular the propagation of excitatory postsynaptic potentials is realistic, and we are easily able to emulate such classical synaptic integration models as direction selectivity. We are also able to emulate the backpropagation into the dendrite of single somatic spikes and bursts of spikes. Thus, this silicon dendrite is suitable for incorporation in detailed silicon neurons operating in real-time; in particular for the emulation of forward- and backpropagating electrical activities found in real neurons.

[1]  C. Koch,et al.  Synaptic background activity influences spatiotemporal integration in single pyramidal cells. , 1991, Proceedings of the National Academy of Sciences of the United States of America.

[2]  M Hines,et al.  A program for simulation of nerve equations with branching geometries. , 1989, International journal of bio-medical computing.

[3]  W. Precht.  The synaptic organization of the brain G.M. Shepherd, Oxford University Press (1975). 364 pp., £3.80 (paperback) , 1976, Neuroscience.

[4]  Bartlett W. Mel.  NMDA-Based Pattern Discrimination in a Modeled Cortical Neuron , 1992, Neural Computation.

[5]  Wilfrid Rall,et al.  Theoretical significance of dendritic trees for neuronal input-output relations , 1964 .

[6]  G Laurent,et al.  A dendritic gain control mechanism in axonless neurons of the locust, Schistocerca americana. , 1993, The Journal of physiology.

[7]  David A. McCormick,et al.  Cellular mechanisms underlying cholinergic and noradrenergic modulation of neuronal firing mode in the cat and guinea pig dorsal lateral geniculate nucleus , 1992, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[8]  Christof Koch,et al.  Biophysics of Computation: Information Processing in Single Neurons (Computational Neuroscience Series) , 1998 .

[9]  David P. M. Northmore,et al.  Building silicon nervous systems with dendritic tree neuromorphs , 1999 .

[10]  Konrad P. Körding,et al.  A learning rule for dynamic recruitment and decorrelation , 2000, Neural Networks.

[11]  Idan Segev,et al.  Compartmental models of complex neurons , 1989 .

[12]  N. Spruston,et al.  Action potential initiation and backpropagation in neurons of the mammalian CNS , 1997, Trends in Neurosciences.

[13]  Misha Mahowald,et al.  Weight vector normalization in an analog VLSI artificial neuron using a backpropagating action potential , 1998 .

[14]  Clay M. Armstrong,et al.  Dendritic Function: Where does it all begin? , 1994, Current Biology.

[15]  M. Livingstone,et al.  Mechanisms of Direction Selectivity in Macaque V1 , 1998, Neuron.

[16]  David P. M. Northmore,et al.  Switched-capacitor neuromorphs with wide-range variable dynamics , 1995, IEEE Trans. Neural Networks.

[17]  W. Rall.  Cable theory for dendritic neurons , 1989 .

[18]  Christoph Rasche.  Analog VLSI circuits for emulating computational features of pyramidal cells , 1999 .

[19]  B. Connors,et al.  Regenerative activity in apical dendrites of pyramidal cells in neocortex. , 1993, Cerebral cortex.

[20]  Carver Mead,et al.  Analog VLSI and neural systems , 1989 .

[21]  J. Brant Arseneau,et al.  VLSI and neural systems , 1990 .

[22]  Christof Koch,et al.  Dynamic range and sensitivity adaptation in a silicon spiking neuron , 1999, IEEE Trans. Neural Networks.

[23]  D. Tank,et al.  Dendritic Integration in Mammalian Neurons, a Century after Cajal , 1996, Neuron.

[24]  Idan Segev,et al.  Excitable dendrites and spines: earlier theoretical insights elucidate recent direct observations , 1998, Trends in Neurosciences.

[25]  B. Sakmann,et al.  A new cellular mechanism for coupling inputs arriving at different cortical layers , 1999, Nature.

[26]  J. C. Anderson,et al.  Dendritic asymmetry cannot account for directional responses of neurons in visual cortex , 1999, Nature Neuroscience.

[27]  H. Barlow.  Vision Science: Photons to Phenomenology by Stephen E. Palmer , 2000, Trends in Cognitive Sciences.

[28]  D. McCormick,et al.  Control of firing mode of corticotectal and corticopontine layer V burst-generating neurons by norepinephrine, acetylcholine, and 1S,3R- ACPD , 1993, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[29]  James M. Bower,et al.  The Book of GENESIS , 1994, Springer New York.

[30]  Rodney J. Douglas,et al.  An Improved Silicon Neuron , 2000 .

[31]  Arthur Konnerth,et al.  Dendritic signal integration , 1997, Current Opinion in Neurobiology.

[32]  John G. Elias,et al.  Artificial Dendritic Trees , 1993, Neural Computation.

[33]  Paul F. M. J. Verschure,et al.  On the Role of Biophysical Properties of Cortical Neurons in Binding and Segmentation of Visual Scenes , 1999, Neural Computation.

[34]  D. Johnston,et al.  Foundations of Cellular Neurophysiology , 1994 .

[35]  Richard H. R. Hahnloser,et al.  Silicon synaptic depression , 2001, Biological Cybernetics.

[36]  R. Douglas,et al.  A silicon neuron , 1991, Nature.

[37]  C. Mead,et al.  Neuromorphic analogue VLSI. , 1995, Annual review of neuroscience.

[38]  Idan Segev.  Single neurone models: oversimple, complex and reduced , 1992, Trends in Neurosciences.

[39]  Ph. Häfliger,et al.  Spike Based Normalizing Hebbian Learning in an Analog VLSI Artificial Neuron , 1999, NIPS 1999.

[40]  Christoph Rasche,et al.  An a VLSI basis for dendritic adaptation , 2001 .

[41]  T. Sejnowski,et al.  Reduced compartmental models of neocortical pyramidal cells , 1993, Journal of Neuroscience Methods.

[42]  Bartlett W. Mel,et al.  Information Processing in Dendritic Trees , 1994, Neural Computation.

[43]  Rodney J. Douglas,et al.  Synchronization of Bursting Action Potential Discharge in a Model Network of Neocortical Neurons , 1991, Neural Computation.