Branching patterns of corticospinal axon arbors in the rodent

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Abstract
Despite extensive study of corticospinal connections in a variety of species, little is known about the detailed morphology of corticospinal axon arbors. Results in previous studies of primates based on intra-axonal filling with horseradish peroxidase (HRP) staining of a limited sample of fibers suggest that corticospinal arbors branch widely to multiple motoneuronal pools. To determine whether this pattern of corticospinal connectivity is present in nonprimate species as well, we studied the branching patterns of corticospinal axon arbors in a rodent species, the golden hamster. The axons were labeled by iontophoretic ijection of Phaseolus vulgaris-leucoagglutinin (PHA-L) into small regions of the forelimb and hindlimb sensorimotor cortex, and immunohistochemistry with the peroxidase-antiperoxidase (PAP) method was used to reveal fine details of terminal arbors within the cervical and lumbar enlargements of the spinal cord. As in higher mammals, corticospinal connections are topographically organized. Moreover, corticospinal axons arising from somatosensory cortx project primarily to the dorsal horn, whereas those from motor cortex terminate most heavily in the ventral horn. This differential projection pattern, not previously demonstrated in rodents, implies functional differences between somatosensory and motor components of the corticospinal pathway. Reconstruction of corticospinal arbors in the ventral horn showed that in both cervical and lumbar spinal cord segments, axons branch widely into interneuronal regions. A surprising number appear to extend into motoneuron cell groups, and some of these axons branch into multiple motoneuronal pools. Widely divergent corticospinal axons that branch to multiple motoneuron pools have been shown to mediate activity in functionally related muscle groups of the primate forearm. The present results suggest that in other species, such as the rodent, a similar divergence of corticospinal arbors may also function to facilitate activity in subsets of muscles.