The Role of Chemical Mechanisms in Neural Computation and Learning

Most computational models of neurons assume that their electrical characteristics are of paramount importance. However, all long-term changes in synaptic efficacy, as well as many short-term effects, are mediated by chemical mechanisms. This technical report explores the interaction between electrical and chemical mechanisms in neural learning and development. Two neural systems that exemplify this interaction are described and modelled. The first is the mechanisms underlying habituation, sensitization, and associative learning in the gill withdrawal reflex circuit in Aplysia, a marine snail. The second is the formation of retinotopic projections in the early visual pathway during embryonic development.

On Directional Selectivity in Vertebrate Retina: An Experimental and Computational Study

This thesis describes an investigation of retinal directional selectivity. We show intracellular (whole-cell patch) recordings in turtle retina which indicate that this computation occurs prior to the ganglion cell, and we describe a pre-ganglionic circuit model to account for this and other findings which places the non-linear spatio-temporal filter at individual, oriented amacrine cell dendrites. The key non-linearity is provided by interactions between excitatory and inhibitory synaptic inputs onto the dendrites, and their distal tips provide directionally selective excitatory outputs onto ganglion cells. Detailed simulations of putative cells support this model, given reasonable parameter constraints. The performance of the model also suggests that this computational substructure may be relevant within the dendritic trees of CNS neurons in general.

Vector-Based Integration of Local and Long-Range Information in Visual Cortex

Integration of inputs by cortical neurons provides the basis for the complex information processing performed in the cerebral cortex. Here, we propose a new analytic framework for understanding integration within cortical neuronal receptive fields. Based on the synaptic organization of cortex, we argue that neuronal integration is a systems--level process better studied in terms of local cortical circuitry than at the level of single neurons, and we present a method for constructing self-contained modules which capture (nonlinear) local circuit interactions. In this framework, receptive field elements naturally have dual (rather than the traditional unitary influence since they drive both excitatory and inhibitory cortical neurons. This vector-based analysis, in contrast to scalarsapproaches, greatly simplifies integration by permitting linear summation of inputs from both "classical" and "extraclassical" receptive field regions. We illustrate this by explaining two complex visual cortical phenomena, which are incompatible with scalar notions of neuronal integration.