Descending pathways to the spinal cord: II. Quantitative study of the tectospinal tract in 23 mammals

To study the early evolution of the mammalian motor systems, we have collected quantitative data on the nuclear origins of tracts descending into the spinal cord in 99 individuals representing 23 species of mammals and one species of reptile. In each individual, the spinal cord was hemisected at the C1-C2 junction and raw HRP immediately applied to the cut fibers. After a 3-day survival period, brain and spinal cord sections were treated with conventional tetramethylbenzidine procedures. In every case, this procedure resulted in heavy retrograde labeling of neural somata throughout the neuraxis from coccygeal cord to cerebral neocortex. Many thousands of supraspinal neurons were vividly labeled within at least 27 discrete cell groups in every mammal (Nudo and Masterton, '88). Despite the vast number and wide diversity of heavily labeled neurons, however, relatively few labeled somata were found in the superior colliculus. The total number of labeled cells in the tectum contralateral to the hemisection was highest in the cat (909) and second highest in the raccoon (628). In the remaining animals, the number was considerably less--averaging only 243 in the 23 mammalian species, 193 in the 21 noncarnivores, and 95 in the iguana. In 7 species of primates the average was 220, and in 3 species of Old World monkeys the average was 142. This wide variation in the number of tectospinal neurons is not related to body size, brain size, or absolute and relative tectum size. Arranging the animals in order of their kinship or recency-of-last-common-ancestor with Man, the average number of labeled tectal cells tends to decrease slightly, whereas arranging the same animals in order of their kinship with the cat or raccoon shows a marked and statistically reliable increase. Neither the evolutionary increase in the tectospinal tract along the Carnivora lineage nor the slight decrease along Man's lineage is altered by mathematical corrections for allometric or scaling factors. Of an array of morphological, visual, motor, and ecological traits tested statistically as a possible source of the variation in size of the tectospinal tract, only a primarily carnivorous feeding preference was found to be reliably related. The relatively small number of tectospinal fibers in most mammals in our sample, including the primates, suggests that the tectospinal tract in Man may be quite small, perhaps far too small to warrant continuing description as a "major descending tract."