Contribution of Cutaneous Inputs From the Hindpaw to the Control of Locomotion. I. Intact Cats

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Abstract
The goal of this study was to evaluate the role of hindpaw cutaneous feedback in the control of locomotion, by cutting some (in one cat) or all (in 2 cats) cutaneous nerves bilaterally at ankle level. Kinematic and electromyographic (EMG) recordings were obtained before and for several weeks after denervation during level and incline (15° up and down) treadmill walking. Ladder walking and ground reaction forces were also documented sporadically. Early after the denervation (1–3 days), cats could not walk across a ladder, although deficits were small during level treadmill walking. Increased knee flexion velocity caused a 14% reduction in swing phase duration. EMG activity was consistently increased in knee, ankle, and toe flexors, and in at least one knee or ankle extensor. The adaptive changes during walking on the incline were much reduced after denervation. Ladder walking gradually recovered within 3–7 wk. By this time, level treadmill walking kinematics had completely returned to normal, but EMG activity in flexors remained above control. Incline walking improved but did not return to normal. Mediolateral ground reaction forces during overground walking were increased by 200%. It is concluded that in intact cats, cutaneous inputs contribute more to demanding situations such as walking on a ladder or on inclines than to level walking. Active adaptive mechanisms are likely involved given that the EMG locomotor pattern never returned to control level. The companion paper shows on the other hand that when the same cats are spinalized, these cutaneous inputs become critical for foot placement during locomotion.