Functional morphology of undulatory pectoral fin locomotion in the stingray taeniura lymma (Chondrichthyes: dasyatidae)
Rajiform locomotion is a unique swimming style found in the batoid fishes (skates and rays) in which thrust is generated by undulatory waves passing down the enlarged pectoral fins. We examined the kinematic patterns of fin motion and the motor patterns of pectoral fin muscles driving the locomotor system in the blue-spot stingray Taeniura lymma. Our goals in this study were to determine overall patterns of fin motion and motor control during undulatory locomotion, to discover how these patterns change with swimming velocity and to correlate muscle function with kinematics and pectoral morphology. Kinematic data were recorded from five individuals over a range of swimming speeds from 22 to 55 cm s(-)(1) (0.9-3.0 DL s(-)(1), where DL is body disc length). Electromyographic (EMG) data were recorded from three individuals over a range of velocities (1.2-3.0 DL s(-)(1)) at seven locations (four dorsal, three ventral) along the pectoral fin. As swimming velocity increases, fin-beat frequency, wavespeed and stride length increase, number of waves and reduced frequency decrease and fin amplitude remains constant. There is variability among individuals in frequency and amplitude at a given speed. An inverse relationship was found in which a high fin-beat frequency is associated with a low fin amplitude and a low fin-beat frequency is associated with a high fin amplitude. The motor pattern of undulatory locomotion is alternate firing activity in the dorsal and ventral muscles as the wave moves along the fin from anterior to posterior. Fin muscles are active along the entire length of the fin except at the lowest speeds. As swimming velocity and fin-beat frequency increase, the time of activation of posterior muscles becomes earlier relative to the onset of activity in the anterior dorsal muscles. The duration of muscle activity is longer in the ventral muscles than in the dorsal muscles, indicating that they play a central role in the power stroke of the fin-beat cycle. The anterior muscles (dorsal and ventral) are active for a relatively longer part of the stride cycle than the posterior muscles. Both the anterior position and the large duty factor of the anterior muscles reflect the role of these muscles in initial wave generation. Synchronous recordings of kinematic data with EMG data reveal that the anterior dorsal and middle ventral muscles do mostly positive work, whereas the dorsal and ventral posterior muscles do negative work at most swimming speeds.