Student Theses and Dissertations

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RU Laboratory

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neuronal variability, nerve impulses, nerve inhibition, eccentric cell, limulus eye


Thesis is concerned with the source and characteristics of variability in the discharge of impulses by neurons. The neuron in which variability was studied is the eccentric cell in the compound eye of the horseshoe crab, Limulus polyphemus. In Part I a theory is presented which accounts for the variability in the response of an eccentric cell to light. The main idea of this theory is that the source of randomness in the impulse rate is "noise" in the generator potential. Another essential aspect of the theory is the view that the process which codes the generator potential into the impulse rate may be treated as a linear filter. These ideas lead directly to Fourier analysis of the fluctuations of the generator potential and fluctuations of the impulse rate. Experimental verification of theoretical predictions was obtained by measurement of the fluctuations and calculation of their variance spectrum. The variance spectrum (or power spectrum) of the impulse rate is shown to be the filtered variance spectrum of the generator potential. Another verification of the theory is the finding that in many cells the signal-to-noise ratio is constant for responses to sinusoidally modulated light, at all modulation frequencies. Inhibition from neighboring eccentric cells will have an effect on the variability of firing of a given eccentric cell. The effects of inhibition are discussed in Part II. The reduction in the average impulse rate which is caused by inhibition decreases the variance of the impulse rate. However, this reduction of the average impulse rate increases the coefficient of variation of the impulse rate. Inhibitory synaptic noise adds to the low frequency portion of the variance spectrum of the impulse rate. This occurs because of the Ill slow time course of the inhibitory synaptic potentials. As a consequence, inhibition decreases the signal-to-noise ratio for low frequency modulated stimuli. The net effect of inhibition is to increase the coefficient of variation of the impulse rate. This effect is predicted by the linear model of the eccentric cell. The same qualitative effect is predicted by other theories of neuronal variability, although its importance is stressed here for the first time.


A thesis presented to the faculty of The Rockefeller University in partial fulfillment of the requirements for the degree of Doctor of Philosophy.

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