Foundations of the Neuron Doctrine

Foundations of the Neuron Doctrine

Foundations of the Neuron Doctrine

Foundations of the Neuron Doctrine

Synopsis

For a century, the neuron doctrine has been the basis for our concepts of nervous organization and brain function. Formulated in 1891 by Wilhelm Waldeyer, it stated that the cell theory applies to the nervous system. Santiago Ramon y Cajal, Spain's greatest scientist, was its main architect; his main tool was a capricious nerve cell stain discovered by Camillo Golgi. This book reviews the original papers on which the neuron doctrine was based, showing that the evidence came from a much wider base of contributions than is generally realized, including such diverse and brilliant personalities as Albrecht Kolliker, Sigmund Freud, Wilhelm His, August Forel, Fritdjof Nansen and Gustav Retzius. Furthermore, many questions about terminology of the parts of the neuron and about the organization of neurons into reflex pathways and networks were raised and debated, questions that remain relevant to this day. Electron microscopical studies in the 1950s appeared to confirm the classical doctrine, but subsequent studies have revealed complexities that were not anticipated. This book reviews these new studies against the background of the classical work, and suggests some new directions for revising our concept of the neuron as a basis for the functional organization of the nervous system.

Excerpt

My interest in this subject began in Oxford in 1959 when my mentors, Charles Phillips and Tom Powell, set me to reading Cajal's account of the organization of the olfactory bulb in preparation for my doctoral work on the electrophysiology of transmission in the olfactory pathway. Immediately intrigued, I committed to memory most of the account of this structure in Cajal "Studies on the Limbic Cortex," originally published in the 1890s and made available in English in Lisbeth Kraft's fine translation in 1955 (London:Lloyd-Luke). Those images, verbal and graphic, have been the inspiration for my view of nervous organization, as they have been for many of my colleagues in neuroscience. I wish to emphasize this, because the analysis and revisions to which I and others have subjected those images in no way detracts from our indebtedness to Cajal; rather, in the true spirit of science, it is the way we pay homage to him.

Eager for more, I purchased Cajal's great two-volume treatise in French on the histology of the nervous system published in 1911. This not only revealed the full breadth and depth of his understanding of nervous organization but also introduced me to the gallery of other distinguished contributors to the early work on the nerve cell. I began to translate some of the classical papers from the 1880s and 1890s, in order to study more closely the original observations and, following Kraft's example, make them available to English-speaking neuroscientists who would otherwise remain unaware of this basic literature of our field.

The next step along the way involved work with Wilfrid Rall, Tom Reese, and Milton Brightman, which revealed that the cells in the olfactory bulb interact in ways not predicted by classical ideas about the organization of neurons as contained in the "Neuron Doctrine" and the "Law of Dynamic Polarization of the Neuron." This again sent me back to the classical authors, to find out exactly what they had meant when they formulated these concepts. I was surprised and delighted to realize that this was a literature abounding in a ferment of ideas about basic principles of nervous organization, with spirited debates over the terminology of the neuron, structure-- function relations at the cellular and circuit level, and rules for the formation of functional pathways and behavioral systems. Several modern authors had begun to reassess the classical doctrine in the light of new find-

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