Thresholds of cortical activation of muslce spindles and alpha motoneurones of the baboon's hand.

1. Much current thinking about voluntary movement assumes that the segmental gamma loops can function as a servomechanism operated by the brain. However, the alpha motoneurones of the baboon's hand receive a powerful monosynaptic (CM) projection from the precentral gyrus. If servo-driving from the same cortical area is to be possible, it must project independently to the fusimotor neurones and have sufficient power to increase the afferent signalling from the muscle spindles. The cortical thresholds for contraction of m. extensor digitorum communis and for acceleration of the discharges of its muscle spindles have therefore been compared.2. Significant results in this context require that the spindles studied be coupled in parallel with the responding extrafusal muscle fibres. Many spindles were not unloaded by the submaximal contractions evoked by cortical stimulation, although all so tested were unloaded by maximal motor nerve twitches. Reasons are given for thinking that such apparent lack of parallel coupling is an artifact of complex intramuscular anatomy and limitation of shortening by ;isometric' myography.3. A brief burst of corticospinal volleys at 500/sec, which is specially effective in exciting alpha motoneurones over the CM projection, failed to excite spindle afferents at or below the threshold for a cortical ;twitch'.4. In a few epileptiform discharges, bursts of spindle acceleration occurred independently of the clonic contractions. A relatively direct and independent cortico-fusimotor (CF) projection may therefore exist.5. Prolonged near-threshold stimulation at 50-100/sec, which allows time for temporal summation in the less direct projections (e.g. cortico-interneuronal, cortico-rubro-spinal) and does not cause frequency-potentiation at CM synapses, gives abundant evidence of independent alpha and fusimotor projections, whose actions hardly outlast the stimulation period.6. Although independent CF projections would permit servo-driving in natural movements of the hand (given adequate loop gain), there has been no evidence of servo-driving by cortical stimulation or in the spontaneous contractions of light anaesthesia.7. Independent projections would provide for controlled alphagamma co-excitation in the servo-governing of natural movements (Matthews, 1964).8. Evidence is reviewed that the CM projection itself may be part of an important control loop for voluntary movement in primates. A corollary would be a diminished importance of CF projections for segmental loops and an increased importance for maintaining the spindle input to cortical loops.