Toe flexor muscle spindle discharge and stretch modulation during locomotor activity in the decerebrate cat

In order to investigate the nature (i.e. static or dynamic) of fusimotor drive to the flexor hallucis longus (FHL) and flexor digitorum longus (FDL) muscles during locomotion we recorded Ia and group II muscle spindle afferent responses to sinusoidal stretch (0.25 and 1 mm amplitude, respectively, 4-5 Hz) in a decerebrate cat preparation. FHL Ia and group II afferents generally had increased discharge rates and decreased modulation to stretch throughout the step cycle, compared to rest, suggesting raised static gamma drive at all locomotor phases. Although the modulation of Ia afferents was reduced during locomotion, most (13 of 18) showed a clear increasing trend during homonymous muscle activity (extension). This was consistent with phasic dynamic gamma drive to FHL spindles linked with alpha drive. In agreement with previous reports, FHL gave a single burst of EMG activity during the step cycle while FDL alpha drive had two components. One was related to extension while the other comprised a brief burst around the end of this phase. Typically FDL Ia and group II afferents also had elevated firing rates and reduced modulation at all locomotor phases, again implicating static gamma drive. Half the afferents (seven Ia, three group II) showed increased discharge during extension, suggesting phasic static gamma drive. There was no gamma drive associated with the late FDL alpha burst. In conclusion, the gamma drives to FHL and FDL differed during locomotion. FHL, which has the alpha drive of a classic extensor, received gamma drive that closely resembled other extensors. The gamma drive of FDL, which exhibits both extensor and flexor alpha synergies, did not match either muscle type. These observations are compatible with the view that fusimotor drive varies in different muscles during locomotion according to the prevailing sensorimotor requirements.