Patterns of fusimotor activity during locomotion in the decerebrate cat deduced from recordings from hindlimb muscle spindles.

1. Recordings have been made from multiple single muscle spindle afferents from medial gastrocnemius (MG) and tibialis anterior (TA) muscles of one hindlimb in decerebrate cats, together with ankle rotation and EMG signals, during treadmill locomotion. Whilst the other three limbs walked freely, the experimental limb was denervated except for the nerves to MG and TA and secured so that it could rotate only at the ankle joint, without any external load. Each afferent was characterised by succinylcholine testing with regard to its intrafusal fibre contacts. Active movements were recorded and then replayed through a servo mechanism to reproduce the muscle length changes passively after using a barbiturate to suppress gamma-motor firing. 2. The difference in secondary afferent firing obtained by subtracting the discharge during passive movements from that during active movements was taken to represent the profile of static fusimotor activity. This indicated an increase before the onset of movement followed by a strongly modulated discharge in parallel with muscle shortening during locomotion. The pattern of static firing matched the pattern of unloaded muscle shortening very closely in the case of TA and with some phase advance in the case of MG. The same effects were observed in primary afferents. 3. Primary afferents with bag1 (b1) contacts in addition showed higher firing frequencies during muscle lengthening in active than in passive movements. This indicated increased dynamic fusimotor firing during active locomotion. There was no evidence as to whether this fluctuated during the movement cycles. 4. When the mean active minus passive difference profile of firing in bag2-chain (b2c) type primary afferents was subtracted from that for b1b2c afferents, the difference was dominated by a peak centred on the moment of maximum lengthening velocity (v). 5. The component of the active minus passive difference firing due to b1 fibre contacts could be modelled by f(t) = av (where a is a constant) during lengthening and by f(t) = 0.2 av during shortening. The remainder of the difference signal matched the predictions of the static fusimotor signal derived from secondary afferents. 6. The findings are discussed in relation to the concept that the modulated static fusimotor pattern may represent a 'temporal template' of the expected movement, though the relationship of the results to locomotion in the intact animal will require further investigation. The analysis of the data indicates that the combined action of muscle length changes and static and dynamic fusimotor activity to determine primary afferent firing can be understood in terms of the interaction between the b1 and b2c impulse initiation sites.