Pattern of motor coordination underlying backward swimming in the lamprey

The main form of locomotion in the lamprey (a lower vertebrate, cyclostome) is forward swimming (FS) based on periodical waves of lateral body flexion propagating from head to tail. The lamprey is also capable of backward swimming (BS). Here we describe the kinematical and electromyographic (EMG) pattern of BS, as well as the effects on this pattern exerted by different lesions of the spinal cord. The BS was evoked by tactile stimulation of a large area in the anterior part of the body. Swimming was attributed to the waves of lateral body undulations propagating from tail to head. The EMG bursts on the two sides alternated, and the EMG in more caudal segments led in phase the EMG in more rostral segments. Main kinematical characteristics of BS strongly differed from those of FS: the amplitude of undulations was much larger and their frequency lower. Also, the maintenance of the dorsal-side-up body orientation ascribed to vestibular postural reflexes (typical for FS) was not observed during BS. A complete transection of the spinal cord did not abolish the generation of forward-propagating waves rostral to the lesion. After a lateral hemisection of the spinal cord, the BS pattern persisted on both sides rostral to the lesion; caudal to the lesion, it was present on the intact side and reduced or abolished on the lesioned side. The role of the spinal cord in generation of different forms of undulatory locomotion (FS and BS) is discussed.