Cat Distal Forelimb Joints and Locomotion: An X-ray Study

The complex construction of the joint apparatus of the cat distal forelimb, which allows the paw three degrees of freedom, poses special requirements on the neural signals controlling the paw position. To understand the electromyography (emg) signals of the distal forelimb muscles during locomotion, it is necessary to know the kinematics of the forelimb joints in detail. As no such information is available, we used the pulsed X-ray technique in trained cats during treadmill locomotion to analyse the angular excursions of the wrist, the metacarpophalangeal (MCP) and the proximal interphalangeal (PIP) joints. X-ray illuminations were done in either the parasagittal or the frontal plane. At the beginning of the stance phase the wrist (WR) and the MCP joints extended slowly, and the PIP joints flexed. Whereas the WR and the PIP joints maintained a constant angular position of approximately 200 degrees and 60 degrees, respectively, throughout the stance phase, extension continued in the MCP joints from 240 degrees at touch-down to 300 degrees at the end of the stance phase. Slightly before lift-off (100 ms) the WR and the MCP joints flexed rapidly. This flexion changed approximately 150 ms after lift-off into a slow extension. The PIP joints extended rapidly at the beginning and at the end of the swing phase, during the interposed period of the swing phase they displayed a slow flexion. Rotatory movements of the forelimb in the radioulnar joints were present during the swing and stance phases. During the swing phase the limb first supinated (starting 100 ms after lift-off); pronation occurred immediately before ground contact. During the stance phase the supination angle was kept constant until 100 ms before lift-off, when a short pronation was found. The paw was kept in an ulnar deviated position throughout the complete step cycle. Ulnar deviation decreased at the end of the swing and stance phases. The results of this study increase our understanding of how the body weight is transmitted on to the ground. They suggest four main functions for the skeletomotor apparatus and the underlying neural commands to secure the forward movement of the animal during the stance phase: (i) preparation and stabilization of a force-transmitting platform; (ii) stabilization of the wrist and the carpal/metacarpal joints; (iii) stabilization of the supination angle; (iv) antigravity control of the extension in the MCP.