Growth of a limb spinal nerve: An ultrastructural study

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Abstract
A quantitative study of the delivery of nerve fibers to the developing limb has been made. Electron microscope (EM) sections of SN 9 in Xenopus were taken at close intervals between dorsal root ganglion and limb-bud. Numbers and types of axons in each section were counted. Counts declined steadily with distance from the ganglion towards the base of the limb. This is largely due to some axons not having grown the full distance at the time of fixation. The slope corresponded with an outgrowth velocity of 0.5 mm a day, without branching. Distally, at stages 50 and 51, but not at stages 49 or 54, the decline was reversed, and a peak of axon number was found. This is strong evidence for axon branching in this region. Since this took place at least one stage before SN 9 bifurcated, and since the bifurcation was distal to the branching when it did appear, it is suggested that axon branching at a nerve bifurcation might be part of a trial-and-error mechanism for pathway selection. Observations were made on the sheath surrounding the nerve, and on its assembly. Initially, axons grow out naked and not parallel; Schwann cells migrate after the growing front. Processes of these cells collect individual axons, and transfer them inwards to form the primitive parallel fibre bundle, enclosed by a family of Schwann cells. Myelination later occurs in a similar manner: Schwann-cell processes extend into a bundle of naked axons and eventually encircle a single transitional axon, with one Schwann cell eventually containing many such axons. Finally comes the stage of promyelin, in which a single Schwann cell encircles a single axon. Myelin whorls are then formed from this complex.