The Role of Constraints in Hebbian Learning

Models of unsupervised, correlation-based (Hebbian) synaptic plasticity are typically unstable: either all synapses grow until each reaches the maximum allowed strength, or all synapses decay to zero strength. A common method of avoiding these outcomes is to use a constraint that conserves or limits the total synaptic strength over a cell. We study the dynamic effects of such constraints. Two methods of enforcing a constraint are distinguished, multiplicative and subtractive. For otherwise linear learning rules, multiplicative enforcement of a constraint results in dynamics that converge to the principal eigenvector of the operator determining unconstrained synaptic development. Subtractive enforcement, in contrast, typically leads to a final state in which almost all synaptic strengths reach either the maximum or minimum allowed value. This final state is often dominated by weight configurations other than the principal eigenvector of the unconstrained operator. Multiplicative enforcement yields a graded receptive field in which most mutually correlated inputs are represented, whereas subtractive enforcement yields a receptive field that is sharpened to a subset of maximally correlated inputs. If two equivalent input populations (e.g., two eyes) innervate a common target, multiplicative enforcement prevents their segregation (ocular dominance segregation) when the two populations are weakly correlated; whereas subtractive enforcement allows segregation under these circumstances. These results may be used to understand constraints both over output cells and over input cells. A variety of rules that can implement constrained dynamics are discussed.

[1]  John H. Holland,et al.  Tests on a cell assembly theory of the action of the brain, using a large digital computer , 1956, IRE Trans. Inf. Theory.

[2]  J. Orbach Principles of Neurodynamics. Perceptrons and the Theory of Brain Mechanisms. , 1962 .

[3]  D. Hubel,et al.  Comparison of the effects of unilateral and bilateral eye closure on cortical unit responses in kittens. , 1965, Journal of neurophysiology.

[4]  R. Guillery Binocular competition in the control of geniculate cell growth , 1972, The Journal of comparative neurology.

[5]  G. Schneider Early lesions of superior colliculus: factors affecting the formation of abnormal retinal projections. , 1973, Brain, behavior and evolution.

[6]  M. Hirsch,et al.  Differential Equations, Dynamical Systems, and Linear Algebra , 1974 .

[7]  R. Pérez,et al.  Development of Specificity in the Cat Visual Cortex , 1975, Journal of mathematical biology.

[8]  D. V. van Essen,et al.  Polyneuronal innervation of skeletal muscle in new‐born rats and its elimination during maturation. , 1976, The Journal of physiology.

[9]  C. Malsburg,et al.  How patterned neural connections can be set up by self-organization , 1976, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[10]  C. Malsburg,et al.  A mechanism for producing continuous neural mappings: ocularity dominance stripes and ordered retino , 1976 .

[11]  T. Sejnowski Statistical constraints on synaptic plasticity. , 1977, Journal of theoretical biology.

[12]  T. Sejnowski,et al.  Storing covariance with nonlinearly interacting neurons , 1977, Journal of mathematical biology.

[13]  Roman Bek,et al.  Discourse on one way in which a quantum-mechanics language on the classical logical base can be built up , 1978, Kybernetika.

[14]  D J Willshaw,et al.  A marker induction mechanism for the establishment of ordered neural mappings: its application to the retinotectal problem. , 1979, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[15]  J. Cowan,et al.  Specificity and plasticity of retinotectal connections: a computational model , 1981, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[16]  E. Oja Simplified neuron model as a principal component analyzer , 1982, Journal of mathematical biology.

[17]  E. Bienenstock,et al.  Theory for the development of neuron selectivity: orientation specificity and binocular interaction in visual cortex , 1982, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[18]  M. Murray,et al.  Target regulation of synaptic number in the compressed retinotectal projection of goldfish , 1982, The Journal of comparative neurology.

[19]  R. Linsker From basic network principles to neural architecture (series) , 1986 .

[20]  R Linsker,et al.  From basic network principles to neural architecture: emergence of spatial-opponent cells. , 1986, Proceedings of the National Academy of Sciences of the United States of America.

[21]  J. Jansen,et al.  Postnatal loss of synaptic terminals in the partially denervated mouse soleus muscle. , 1987, Acta physiologica Scandinavica.

[22]  B. Gustafsson,et al.  Long-term potentiation in the hippocampus using depolarizing current pulses as the conditioning stimulus to single volley synaptic potentials , 1987, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[23]  T. Teyler,et al.  Long-term potentiation. , 1987, Annual review of neuroscience.

[24]  R. L. Meyer,et al.  Retinotopically inappropriate synapses of subnormal density formed by surgically misdirected optic fibers in goldfish tectum. , 1988, Brain research.

[25]  R. L. Meyer,et al.  Optic synapse number but not density is constrained during regeneration onto surgically halved tectum in goldfish: HRP‐EM evidence that optic fibers compete for fixed numbers of postsynaptic sites on the tectum , 1988, The Journal of comparative neurology.

[26]  J. Lisman,et al.  Feasibility of long-term storage of graded information by the Ca2+/calmodulin-dependent protein kinase molecules of the postsynaptic density. , 1988, Proceedings of the National Academy of Sciences of the United States of America.

[27]  Teuvo Kohonen,et al.  Self-organization and associative memory: 3rd edition , 1989 .

[28]  R. L. Meyer,et al.  Impulse blockade by intraocular tetrodotoxin during optic regeneration in goldfish: HRP-EM evidence that the formation of normal numbers of optic synapses and the elimination of exuberant optic fibers is activity independent , 1989, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[29]  K. Miller,et al.  Ocular dominance column development: analysis and simulation. , 1989, Science.

[30]  R. L. Meyer,et al.  Normal numbers of retinotectal synapses during the activity-sensitive period of optic regeneration in goldfish: HRP-EM evidence implicating synapse rearrangement and collateral elimination during map refinement , 1989, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[31]  P. Rakić,et al.  Synaptogenesis in visual cortex of normal and preterm monkeys: evidence for intrinsic regulation of synaptic overproduction. , 1989, Proceedings of the National Academy of Sciences of the United States of America.

[32]  Kenneth D. Miller,et al.  Derivation of Linear Hebbian Equations from a Nonlinear Hebbian Model of Synaptic Plasticity , 1990, Neural Computation.

[33]  E. Knudsen,et al.  Sensitive and critical periods for visual calibration of sound localization by barn owls , 1990, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[34]  D. Mackay,et al.  Analysis of Linsker's application of Hebbian rules to linear networks , 1990 .

[35]  David J. C. MacKay,et al.  Analysis of Linsker's Simulations of Hebbian Rules , 1990, Neural Computation.

[36]  D. V. van Essen,et al.  Synaptic dynamics at the neuromuscular junction: mechanisms and models. , 1990, Journal of neurobiology.

[37]  Michael Merzenich,et al.  Hebb-Type Dynamics is Sufficient to Account for the Inverse Magnification Rule in Cortical Somatotopy , 1990, Neural Computation.

[38]  B. Finlay,et al.  Compensation for population size mismatches in the hamster retinotectal system: Alterations in the organization of retinal projections , 1991, Visual Neuroscience.

[39]  Graeme Mitchison,et al.  Removing Time Variation with the Anti-Hebbian Differential Synapse , 1991, Neural Computation.

[40]  J. Kaas Plasticity of sensory and motor maps in adult mammals. , 1991, Annual review of neuroscience.

[41]  X. D. Yang,et al.  Initial synaptic efficacy influences induction and expression of long-term changes in transmission. , 1991, Proceedings of the National Academy of Sciences of the United States of America.

[42]  K. Miller Development of orientation columns via competition between ON- and OFF-center inputs. , 1992, Neuroreport.

[43]  Nathan Intrator,et al.  Objective function formulation of the BCM theory of visual cortical plasticity: Statistical connections, stability conditions , 1992, Neural Networks.

[44]  R. Malenka,et al.  The influence of prior synaptic activity on the induction of long-term potentiation. , 1992, Science.

[45]  Harry G. Barrow,et al.  The Role of Weight Normalization in Competitive Learning , 1994, Neural Computation.

[46]  Yong Liu,et al.  Influence Function Analysis of PCA and BCM Learning , 1994, Neural Computation.

[47]  L F Abbott,et al.  Decoding neuronal firing and modelling neural networks , 1994, Quarterly Reviews of Biophysics.

[48]  Barak A. Pearlmutter Time-Skew Hebb Rule in a Nonisopotential Neuron , 1995, Neural Computation.

[49]  Colin Fyfe,et al.  Introducing Asymmetry into Interneuron Learning , 1995, Neural Computation.

[50]  Marco Idiart,et al.  Reduced Representation by Neural Networks with Restricted Receptive Fields , 1995, Neural Computation.