Dorsal root projections in the clawed toad (Xenopus laevis) as demonstrated by anterograde labeling with horseradish peroxidase

Abstract

Horseradish peroxidase was applied to the proximal stumps of severed cervical, thoracic and lumbar dorsal roots in the clawed toad, Xenopus laevis, in order to study the course, distribution and site of termination of dorsal root fibers in the spinal cord and brain stem. The anterograde transport of horseradish peroxidase as applied in the present study proved to be a useful and reliable technique. Results show that on entering the spinal cord, dorsal root fibers segregate into a medially placed component entering the dorsal funiculus and a more laterally situated bundle in the dorsal part of the lateral funiculus. As regards its position the latter bundle presumably represents the anuran homologue of the mammalian tract of Lissauer. Moreover, a small intermediate bundle of fibers directly enters the spinal gray matter. The labeled fibers entering the dorsal funiculus and the tract of Lissauer ascend and descend in the spinal cord, displaying a longitudinal arrangement resembling that of higher vertebrates. In the spinal gray, dorsal root fibers terminate in the dorsal, central and lateral fields of Ebbesson, with the last field being a major terminus for dorsal root fibers originating in the limb-innervating segments. No dorsal root fibers were found to project to the motoneuron fields. A dorsal column nucleus, which is divisible into medial and lateral compartments, is present in the obex region and extends from the level of the second spinal nerve to that of the entrance of the vagus and glossopharyngeal nerves. Dorsal root fibers from the lumbar and all thoracic segments project to the medial compartment of the dorsal column nucleus, whereas those of the cervical enlargement project to the lateral compartment. Although the anuran dorsal column nucleus appears to be less differentiated than that of higher vertebrates, its medial and lateral compartments can be considered to be the forerunners of the mammalian nucleus gracilis and nucleus cuneatus, respectively.