Origin of ascending and spinal pathways from the nucleus tegmenti pedunculopontinus in the rat

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Abstract
The distribution and collaterlization of ascending and descending projections from neurons in the nucleus tegmenti pedunculopontinus (PPN) were studied in the rat by using retrograde transport of HRP, HRP/WGA, and fluorescent dyes. The PPN and its two subdivisions, the subnucleus compactus (PPNc) and subnucleus dissipatus (PPNd), were delineated on sagittal Nisslstained sections by using cytoarchitectural features as guidelines. Large bilateral pressure injections of HRP and/or fluorescent dyes into the cervical cord retrogradely labeled moderate numbers of fusiform and polygonal PPN cells which ranged in size between 65 and 390 μm2. The labeled cells were scattered throughout the PPNd and were somewhat more numerous in the medial half of the subnucleus. The PPNc contained only occasional labeled cells in its ventralmost portion. Following single unilateral HRP/WGA injections in the striatum, globus pallidus, entopeduncular nucleus, subthalamus, or the substantia nigra, the distribution of the labeled cells was similar to that of the spinal cord-projecting PPN neurons. Multiple HRP injections were then made bilaterally in the substantia nigra and the entopeduncular nucleus and/or subthalamus in order to label the entire population of PPN neurons projecting to the basal ganglia. In this case, not only the PPNd but also the PPNc contained a substantial number of retrogradely labeled cells. The rostrally projecting PPN cells outnumbered 5.4 times those projecting to the spinal cord, and their somata were somewhat larger, ranging between 114 and 472 μm2. While both fusiform and polygonal shapes were encountered, the polygonal cell somata were more numerous. In the double-labeling experiments, Granular Blue and Diamidino Yellow Dihydrochloride were injected into the cervical cord and the entopeduncular nucleus or subthalamus. In general, these experiments confirmed the extensive overlap of forebrain- and spinal cord-projecting neurons within the PPNd and the quantitative preponderance of ascending neurons. They also demonstrated that these two projection systems originate largely from separate cell populations since the double-labeled cells always composed less than 5% of the labeled neurons. The results confirm the existence of a direct PPN projection to the spinal cord. This pathway originates mainly in the PPNd and appears to be quantitatively weaker than the PPN projections to the forebrain. The spinal cord-projecting cells are not spatially segregated from the cells projecting to the basal ganglia, but they represent a separate population of the PPN projection neurons.