Different forms of locomotion in the spinal lamprey

Forward locomotion has been extensively studied in different vertebrate animals, and the principal role of spinal mechanisms in the generation of this form of locomotion has been demonstrated. Vertebrate animals, however, are capable of other forms of locomotion, such as backward walking and swimming, sideward walking, and crawling. Do the spinal mechanisms play a principal role in the generation of these forms of locomotion? We addressed this question in lampreys, which are capable of five different forms of locomotion - fast forward swimming, slow forward swimming, backward swimming, forward crawling, and backward crawling. To induce locomotion in lampreys spinalised at the second gill level, we used either electrical stimulation of the spinal cord at different rostrocaudal levels, or tactile stimulation of specific cutaneous receptive fields from which a given form of locomotion could be evoked in intact lampreys. We found that any of the five forms of locomotion could be evoked in the spinal lamprey by electrical stimulation of the spinal cord, and some of them by tactile stimulation. These results suggest that spinal mechanisms in the lamprey, in the absence of phasic supraspinal commands, are capable of generating the basic pattern for all five forms of locomotion observed in intact lampreys. In spinal lampreys, the direction of swimming did not depend on the site of spinal cord stimulation, but on the stimulation strength. The direction of crawling strongly depended on the body configuration. The spinal structures presumably activated by spinal cord stimulation and causing different forms of locomotion are discussed.