RESPONSES OF CAT MUSCLE-SPINDLES WHICH LACK A DYNAMIC FUSIMOTOR SUPPLY

1. The experiments reported here support the view that some spindles in the peroneus tertius muscle of the anaesthetized cat lack a nuclear bag1 intrafusal fibre. 2. The bag1 fibre is characterized by the fact that it is innervated exclusively by dynamic fusimotor axons. A method was devised to test each spindle in peroneus tertius for a dynamic fusimotor innervation. The ventral roots containing the muscle's motor supply were subdivided into five portions, approximately equal in terms of the tension they generated, and each piece was stimulated in turn, repetitively, at fusimotor strength, during ramp stretch of the muscle, to look for a large increase in dynamic response. 3. The method allowed confirmation that the majority of spindles in peroneus tertius had a dynamic fusimotor innervation. However, where the dynamic effect was weak and accompanied by a strong static fusimotor action and extrafusal unloading, it risked being overlooked. 4. The confirmatory test for the presence of a bag1 fibre was whether or not the spindle showed a large increase in dynamic response in the presence of the drug succinyl choline (SCh) injected arterially close to the muscle in which the spindle is located. SCh is known to induce a contracture in the bag1 fibre and therefore mimics tonic dynamic fusimotor stimulation. 5. In five experiments, of a total of forty-two spindles with afferents conducting within the group I range, five examples were encountered where there was no increase in dynamic response, either with ventral root stimulation or perfusion with SCh. It was concluded that these were spindles which lacked a bag1 fibre. 6. Passive stretch of such spindles revealed no feature in the response which allowed them to be distinguished from spindles in which the bag1 fibre was present. This conclusion posed the question, what contribution, if any, does the bag1 fibre make to the stretch response? 7. It was possible to show that under some conditions the bag1 fibre did contribute to the response to stretch. If the spindle was conditioned by repetitive nerve stimulation, at fusimotor strength, at a length longer than that at which a test stretch was applied, the response to the stretch was delayed, and it began at a lower rate than after conditioning stimulation at the test length. The delayed response was attributed to the presence of slack in intrafusal fibres. With the spindle in this state, if a second conditioning stimulus was given to a single dynamic fusimotor fibre, supplying the spindle, the onset of the stretch response was earlier, suggesting that slack had been taken up selectively in the bag1 fibre, which was now able to contribute to the stretch response. 8. It is concluded that during large ramp stretches of passive spindles the dynamic response is normally generated in the bag2 fibre. The bag1 fibre makes a significant contribution only if it is actively contracting or when the stretch immediately follows a period of dynamic fusimotor stimulation.