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The anterograde tracer Phaseolus vulgaris-leucoagglutinin (PHA-L) was used to study the distribution and density of the projections that originate from four identified subdivisions of the pericruciate cortex (namely, the forelimb and hind limb representations of area 4, area 6a beta, and area 6a gamma) and that terminate in the pontomedullary brainstem in the cat. Injections of PHA-L in all areas of the pericruciate cortex labelled numerous fibers and their terminal swellings in the brainstem. The major target regions of all four cortical areas were the pontine nuclei and the pontomedullary reticular formation (PMRF). Injections into both the forelimb and hind limb representations of area 4 and into area 6a beta resulted in a dense pattern of terminal labelling in restricted regions of the medial and lateral parts of the ipsilateral pontine nuclei. The labelling following the area 6a beta injection was spatially distinct from that seen following the area 4 injections. Injections into the forelimb representation of area 4 as well as into area 6a beta and 6a gamma resulted in the labelling of numerous terminal swellings bilaterally in the PMRF; in contrast, there were few labelled terminal swellings in the PMRF following injections into the hind limb representation of area 4. Terminal swellings on individual corticoreticular fibers were far less densely aggregated than those in the pontine nuclei. The dense pattern of innervation to restricted regions of the pontine nuclei supports previous suggestions that the corticopontine projections retain a high degree of topographical specificity that could be used in the control of discrete voluntary movements. In contrast, the more diffuse pattern of the projections to the PMRF may facilitate the selection and activation of the complex postural patterns that accompany voluntary movement.

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The purpose of this study was 1). to characterize the morphology of lumbar commissural neurons (CNs) with reticulospinal inputs and 2). to quantitate their activity during locomotor rhythm generation. Intraaxonal recordings at the L4-7 level of the spinal cord were obtained in 67 neurons in the decerebrate, paralyzed cat. Fourteen of them were subsequently nearly fully visualized following their intraaxonal injection with the tracer neurobiotin. All 14 were CNs with axons projecting across the midline of the spinal cord. Their somata were located mainly in lamina VIII and additionally in laminae VII-VI. Most of the lamina VIII CNs were excited monosynaptically from reticulospinal pathways. They were judged to be interneuronal CNs if they had no, or a short, rostral projection. These CNs commonly gave off multiple axon collaterals in and around their somata's segmental level. They projected mainly to laminae VIII-VII and some additionally to lamina IX. Some laminae VIII and the laminae VII-VI CNs were excited polysynaptically from reticulospinal pathways or were not excited. They were judged to be long propriospinal or ascending tract CNs because they had only an ascending axon. Most lamina VIII CNs discharged rhythmically during fictive locomotion evoked by stimulation of the mesencephalic locomotor region, exhibiting one peak per locomotor cycle. The peak was in phase with neurographic activity of either a left or a right hindlimb extensor nerve. These results suggested that lamina VIII CNs are reciprocally connected bilaterally at each segmental level. Such an arrangement suggests their participation in the generation and coordination of reciprocal and bilateral locomotor activity.