Abstract
We studied the responses of 34 deefferented spindle receptors to slowly applied ramp stretches (0.01-1 mm/s) of small (0.02-0.2 mm) and intermediate (0.2-1 mm) amplitudes. The afferent discharge from primary and secondary endings was recorded from filaments of dorsal root in anesthetized cats. 1. Responses of most endings to ramps of intermediate amplitude showed abrupt changes in slope (discontinuities) which were highly repeatable. Discontinuities occurred more nearly at constant stretch (in the range 50-400 mum for different receptors) than at constant discharge rate. They were less pronounced in the case of secondary endings. 2. Changes in sensitivity occurred when the degree of stretch exceeded a transitional amplitude which ranged from 50 to 200 mum. These changes were studied by constructing plots based on a family of responses to a family of ramps which were scaled versions of each other. The plots indicated that reductions in sensitivity occurred both during stretch and during adaptation; the reductions were more marked for primary than for secondary endings. 3. Responses were modified considerably by preceding changes in muscle length. When the last change was an increase of a few millimeters, discontinuities became more pronounced and other changes in the appearance of the dynamic response occurred, particularly in the case of primary endings. These changes could last for several minutes, but were abolished by a single test stretch of intermediate amplitude. 4. The resetting of high sensitivity that occurs when muscle length is changed, the discontinuities, the transitions in sensitivity, nonlinear adaptation, and the effects of previous length change appeared to be related phenomena. They can all be accounted for by the hypothesis that polar zones of intrafusal muscle fibers possess a frictionlike property, one analogous to that which has been described for whole muscle. A simple nonlinear model which shows these features is presented. 5. The adequate stimulus for a change in primary ending discharge is a small change in muscle length, relatively independently of its velocity. The dynamic response arises mainly from a changing sensitivity to length itself, which is a nonlinear property.