Swimming in the rat: Analysis of locomotor performance in comparison to stepping

Summary

Swimming in a mammalian quadruped, the rat, is analyzed in kinematic (joint angles) and electromyographic (EMG) terms. Data were collected on the movements of the hip, knee, ankle, and toe joints and three principle extensors and three flexors of the right hindlimb and compared with similar data collected on the same rats during treadmill stepping. The flexion, or protraction phase of swimming and stepping had many elements in common, including a similarity of EMG activity patterns and corresponding limb movements. However, in the extension, or retraction phase, there were notable differences. Although joint-extensor muscles were all coactive in both conditions, the brevity of the swimming extensor phase precluded the characteristic variation in EMG activity levels seen in the extensors in stepping. The flexors, in particular semitendinosus (ST), exhibited bursts of activity at the end of the extensor phase of swimming which were not present during the comparable period of stepping. The extra burst in ST produced a very rapid knee flexion at this time. Whereas the range of hip joint movement was similar in the two conditions, the ranges of the knee and ankle joints were expanded during swimming.

Overall, the evidence suggests that swimming is a very rapid form of a basic locomotor pattern in which the extensors are driven to their maximum contraction rate. The extra extension of the limb derives from the absence of ground reaction forces, allowing the knee and ankle joints to fully extend. The added bursts in the flexors remain to be explained. A discussion of these results in terms of current theories of single limb locomotor pattern generation is presented.