Adaptive control for backward quadrupedal walking. I. Posture and hindlimb kinematics

Abstract

1. To gain new perspectives on the neural control of different forms of quadruped locomotion, we studied adaptations in posture and hindlimb kinematics for backward (BWD) walking in normal cats. Data from four animals were obtained from high-speed (100 fr/s) cine film of BWD treadmill walking over a range of slow walking speeds (0.3-0.6 m/s) and forward (FWD) treadmill walking at 0.6 m/s. 2. Postural adaptations during BWD walking included flexion of the lumbar spine, compared to a relatively straight spine during FWD walking. The usual paw-contact sequence for FWD walking [right hindlimb (RH), right forelimb (RF), left hindlimb (LH), left forelimb (LF)] was typically reversed for BWD walking (RH, LF, LH, RF). The hindlimbs alternated consistently with a phase difference averaging 0.5 for both forms of walking, but the phasing of the forelimbs was variable during BWD walking. 3. As BWD walking speed increased from 0.3 to 0.6 m/s, average hindlimb cycle period decreased 21%, stance-phase duration decreased 29%, and stride length increased 38%. Compared to FWD walking at 0.6 m/s, stride length was 30% shorter, whereas cycle period and stance-phase duration were 17% shorter for BWD walking. For both directions, stance occupied 64 +/- 4% (mean +/- SD) of the step cycle. 4. During swing for both forms of walking, the hip, knee, and ankle joints had flexion (F) and extension (E1) phases; however, the F-E1 reversals occurred earlier at the hip and later at the knee for BWD than for FWD walking. At the ankle joint, the ranges of motion during the F and E1 phases were similar for both directions. During BWD walking, however, the knee flexed more and extended less, whereas the hip flexed less and extended more. Thus horizontal displacement of the limb resulted primarily from hip extension and knee flexion during BWD swing, but hip flexion and knee extension during FWD swing. 5. At the knee and ankle joints, there were yield (E2) and extension (E3) phases during stance for both forms of walking; however, yields at the knee and ankle joints were reduced during BWD walking. At the hip, angular motion was unidirectional, as the hip flexed during BWD stance but extended during FWD stance. Knee extension was the prime contributor to horizontal displacement of the body during BWD stance, but hip extension was the prime contributor to horizontal displacement during FWD stance. 6. Our kinematic data revealed two discriminators between BWD and FWD walking.(ABSTRACT TRUNCATED AT 400 WORDS)