Organization of the motor nuclei in the cervical spinal cord of salamanders

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Abstract
The distribution and cytoarchitecture of motor nuclei of the cervical spinal cord were studied by using HRP techniques (whole mounts and sections) in 22 species of salamanders (families Hynobiidae, Dicamptodontidae, Ambystomatidae, Salamandridae, and Plethodontidae) representing a wide variety of life histories and functional modes of feeding. The nucleus of the first spinal nerve extends from the level of, or slightly caudad to, the root of the tenth cranial nerve, almost to the ventral root of the second spinal nerve. Approximately one‐half of this nucleus is situated in the brainstem. This anterior extension is longest in bolitoglossine plethodontids. The nucleus of the second spinal nerve extends from the root of the first spinal nerve to the dorsal root of the second spinal nerve. The nuclei of the first and second spinal nerves in all species except bolitoglossines have motor neurons arranged in two columns: a lateral one containing large spindle‐shaped cells and a medial one containing pearshaped or polygonal smaller cells. The primary dendrites of these lateral and medial cells are parallel and their arborization is relatively narrow. In contrast, bolitoglossines lack the lateral motor column. The nucleus of the first spinal nerve consists only of a medial band of pear‐shaped and sometimes polygonal cells, and the nucleus of the second spinal nerve is a wider band of pear‐shaped and polygonal cells which are always situated inside the periventricular gray matter. The arrangement of the somata in bolitoglossines is less organized and the primary dendrites are less parallel and have a broader arborization than in other salamanders. In all species, cells in the second spinal nucleus are arranged in a less orderly manner than those in the first. All salamanders studied possess a spinal accessory nerve whose motor neurons are located in the cervical spinal cord; the axons leave the brainstem with fibers of the vagus nerve. The rostrocaudal extent of this nucleus differs markedly among species. In bolitoglossines the nucleus is more or less restricted to the region of the nucleus of the second spinal nerve. In all other species studied, the accessory nucleus extends from the obex to the caudal end of the nucleus of the third spinal nerve. In the tribe Plethodontini the cytoarchitecture of the accessory nucleus is similar to that of the second spinal. In desmognathine and hemidactyliine plethodontids as well as in all nonplethodontid species studied the nucleus consists of pear‐shaped and cone‐shaped cells. No separate lateral motor column is present and the cone‐shaped cells are found at various positions inside and slightly outside the gray matter. In bolitoglossines both pear‐shaped and polygonal cells are found in the gray matter; again, no lateral motor column is present. A developmental‐evolutionary hypothesis is presented to account for the unique pattern of cellular organization in the bolitoglossines, and functional and phylogenetic implications of our findings are considered.