Collateralization of descending pathways from the brainstem to the spinal cord in a lizard, Varanus exanthematicus

Abstract

With the multiple fluorescent retrograde tracer technique, the collateralization in the spinal cord of descending supraspinal pathways was studied in a lizard, Varanus exanthematicus. Fast Blue (FB) gels were implanted unilaterally in the spinal gray matter of the cervical enlargement and Nuclear Yellow (NY) gels were implanted ipsilaterally in two series of experiments in all spinal funiculi of the lumbar enlargement or in midthoracic spinal segments, respectively. All brainstem nuclei known to project to the spinal cord in reptiles were found to give rise to branching axons that may influence widely separate levels of the spinal cord. The number of double-labeled FB-NY cells varied in these brainstem nuclei from none to half the number of neurons projecting to the cervical enlargement. Highly collateralizing projections (expressed as the percentage of double-labeled neurons, DL) were found to arise from the nucleus raphes inferior, the contralateral nucleus reticularis superior pars lateralis, the contralateral nuclei vestibulares ventromedialis and descendens, and the ipsilateral nucleus reticularis inferior pars ventralis. A lower percentage of DL neurons was noted for the contralateral nucleus ruber and bilaterally for the nucleus reticularis medius and nucleus reticularis inferior. Extensive brainstem projections directed to cervical and high thoracic spinal levels originate from the area lateralis hypothalami, the nucleus of the fasciculus longitudinalis medialis, the contralateral nucleus cerebellaris medialis, and from the nucleus tractus solitarii. Projections preferentially directed to midthoracic or lower levels of the spinal cord were found to arise from the ipsilateral locus coeruleus, the contralateral nucleus reticularis superior pars lateralis, the nucleus reticularis inferior pars ventralis, the nucleus reticularis inferior, and the nucleus raphes inferior. In contrast to findings in mammals, in Varanus exanthematicus the red nucleus, the nucleus vestibularis ventrolateralis, and certain parts of the reticular formation did not display a clear-cut somatotopic organization. In general two different patterns of collateralization can grossly be discerned: a gradual decrease of spinal collaterals caudalward, which can be interpreted as a certain specificity of such projections; and a constant number of collateral nerve fibers throughout the spinal cord that can be interpreted as either a nonspecific or, in contrast, a highly specific system, focussed exclusively on the cervical and lumbar enlargements.