Chapter 2 Comparative anatomy and physiology of the corticospinal system

The corticospinal tract provides the most direct pathway over which the cerebral cortex controls movement. In rodents and marsupials this influence is exerted largely upon interneurons in the dorsal horn of the spinal gray matter. However, ascending the phylogenetic scale through carnivores and primates, the number of corticospinal axons grows and corticospinal terminations shift progressively toward the interneurons of the intermediate zone and ventral horn, ultimately forming increasing numbers of synaptic terminations directly on the motoneurons themselves. Based on this phylogenetic trend, humans are believed to have more direct corticomotoneuronal synapses than any other species, consistent with observations that humans suffer more extensive loss of motility from lesions of the corticospinal tract than do other mammals. Beyond this phylogenetic trend, studies of the corticospinal system in animals have provided insight into the motor abnormalities that result from corticospinal lesions in humans. Corticospinal lesions impair many functionally related muscles and movements in parallel, both because of the divergent output from single corticomotoneuronal cells to multiple motoneuron pools, and because of the convergent input to different motoneuron pools from large, overlapping cortical territories. Furthermore, the weakness, slowness and inflexible, stereotyped movements that remain after corticospinal lesions reflect the loss of input to spinal interneurons and motoneurons from corticospinal neurons, the discharge frequency of which varies with the force, direction and speed of both gross and fine movements. That these deficits resulting from corticospinal lesions are more prominent in humans than in animals indicates, moreover, that animals make greater use of additional descending pathways to control movement. Animal studies have shown that although the bulk of the corticospinal tract arises from the primary motor cortex, this projection is not the only route via which the brain controls movement. Adjacent areas in the frontal and parietal lobes also contribute axons to the corticospinal tract, as well as having corticocortical connections with the motor cortex. Furthermore, the motor cortex and premotor cortex both project to the red nucleus and to the pontomedullary reticular formation, from which the rubrospinal and reticulospinal tracts arise. However, given the limitations on experimental studies in humans, comparative animal studies of the distributed descending system through which the brain controls movement continue to provide deeper understanding and insight into the deficits resulting from human corticospinal lesions, whether caused by stroke, tumor, multiple sclerosis, trauma or ALS.