Light and electron microscopic evidence for a direct corticospinal projection to superficial laminae of the dorsal horn in cats and monkeys.

The anterograde transport of horseradish peroxidase (HRP) and wheat germ agglutin conjugated to horseradish peroxidase (WGA-HRP) was employed in cats and monkeys to investigate, at both the light and electron microscopical levels, the contribution of the corticospinal tract (CST) to the superficial laminae of the dorsal horn. At the light microscopic level, this approach not only confirmed the previously documented pattern of CST termination, but also revealed a sparse projection to laminae VIII and IX of the cat and a prominent projection to the most superficial parts of the brachial dorsal horn, i.e., laminae I and II. Discrete injections involving particular cytoarchitectonic areas (4, 3a, 3b, and 1-2) of monkeys showed that the superficial laminae receive their corticofugal inputs primarily from areas 3b, 1, and 2. Electron microscopic observations were made on CST fibers and boutons which were labelled, after histochemical processing, with the reaction product of anterogradely transported WGA-HRP. The labelled fibers in the superficial laminae were small (+/- 0.5 micron), and boutons established mainly axodendritic contacts, contained mostly clear, spherical, or pleomorphic vesicles, but sometimes also displayed dense core vesicles. These boutons were primarily in lamina I and outer lamina II, but not in inner lamina II. The possible role of a direct monosynaptic pathway from the cerebral cortex to the superficial laminae of the dorsal horn is discussed in relation to the previous reports that laminae I and II play a significant role in nociception.