Retinal ganglion cells : physiology and prosthesis
Abstract
The retina is responsible for encoding different aspects of the visual world. Light enters the eyes and is converted by the photoreceptors into electrochemical signals. These signals are processed by the retinal network and proceed afferently to the brain via the axons of the retinal ganglion cells (RGCs). The RGCs outputs are in the form of action potentials (spikes), which encrypt the visual information in terms of spike shape, firing frequencies, and the firing patterns. When the photoreceptors are gone due to disease, vision is lost. The idea of a retinal prosthesis is to activate the surviving RGCs by electrical stimulation in order to recreate vision. In this thesis, I have studied the physiological properties of the RGCs, and reconstructed natural RGC spike trains by electrical stimulation. Chapter 1 introduces the anatomy of the retina and the retinal neurons. How the RGCs respond to light. Electrical stimulation is also discussed. A brief historical summary of the receptive field properties and cell physiology is also presented. Chapter 2 characterizes the intrinsic properties of 16 morphologically defined types of rat RGCs. The intrinsic properties include the biophysical properties due to morphology and dendritic stratification, in addition to physiological properties such as firing behaviours. These properties are also compared with the cat RGC intrinsic properties in order to investigate the variations between the morphologically similar RGCs of the two species. The results suggest that the RGCs among species, even with similar morphologies, do not have conservative intrinsic properties. Chapter 3 examines the details of the spiking properties of the different rat RGC types. Spikes are initiated at the axonal initial segment. A 'single' spike recorded at the soma consists of an axonal spike and a somatic spike. The existence of the two spikes can be recognized by two humps in the phase plot, and further revealed in the higher derivatives of the membrane potential. A principal component analysis shows that the parameters extracted from the phase plots are very useful for a model-independent rat RGC classification. Chapter 4 establishes the foundations for electrical stimulation of the retina. The question is to what extent optimum placement of the stimulating and reference electrodes might be affected by anatomical location. Here we placed the stimulating electrode above or below the retinal inner limiting membrane and found no statistical difference between the thresholds. In addition, reflective axonal spikes from the cut end are discussed. Chapter 5 combines the knowledge obtained in the previous chapters for the sole purpose of reproducing natural RGC outputs when using electrical stimulation. The light responses of the eye under saccadic movements were recorded and used to form the stimulus patterns. The reconstructions were performed on the brisk-transient (BT) and the brisk-sustained (BS) RGCs. Our results suggested that BT RGCs are more capable of following the stimulated stimulus patterns over a wide range of frequencies than the BS RGCs. Chapter 6 concludes the whole thesis.
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