AbstractThe origins of the descending spinal pathways in sea lampreys (Petromyzon marinus), silver lampreys (Ichthyomyzon unicuspis), and Pacific hagfish (Eptatretus stouti) were identified by the retrograde transport of horseradish peroxidase (HRP) placed in the rostral spinal cord. In lampreys, the majority of HRP‐labeled cells were located along the length of the brainstem reticular formation in the inferior, middle, and superior reticular nuclei of the medulla, mesencephalic tegmentum, and nucleus of the medial longitudinal fasciculus. Labeled reticular cells included the Mauthner and Müller cells. Horseradishperoxidase‐filled cells were also present in the descending trigeminal tract, intermediate and posterior octavomotor nuclei, and a diencephalic cell group, the nucleus of the posterior tubercle. As in lampreys, the reticular formation of the Pacific hagfish was the largest source of descending afferents to the spinal cord. Labeled cells were found in the dorsolateral and ventromedial reticular nuclei, the dorsal tegmentum at the juncture of the medulla and midbrain, and the nucleus of the medial longitudinal fasciculus. Additional medullary cells projecting to the cord were located in the perivagal nucleus, the central gray, and the anterior and posterior magnocellular octavolateralis nuclei. The existence of reticulospinal and possible vestibulo‐, trigemino‐, and solitary spinal projections in lampreys and hagfishes and the wide distribution of these pathways in jawed vertebrates suggest that they evolved in the common ancestor of gnathostomes and both groups of jawless fishes. However, descending spinal pathways from the cerebellum, red nucleus, and telencephalon appear to be gnathostome characters.