A continuing controversy surrounds the question of whether Golgi tendon organs are examples of receptors in which impulses may be generated at more than one site. This paper reports a systematic examination of a number of models incorporating single or multiple impulse generators and of the compatibility of their predictions with experimental observations. Two phenomena, in particular, that must be accounted for are nonlinear summation and cross-adaptation. When two motor units each with a direct effect on the tendon organ are stimulated together, the rate of discharge is greater than either individual rate but is less than their sum. In cross-adaptation a conditioning response elicited by one motor unit contraction produces adaptation of the discharge associated with stimulation of a second motor unit. A model with a central impulse generator can be modified to account for nonlinear summation by postulating a nonlinear transformation in the generator current-to-impulse rate conversion. Experiments measuring summation of responses to stimulation of three inputs produced results that did not support this model. Another variation of the model, which had a nonlinearity in the tension-to-current step and cross-connections (mechanical or neural) between tendon strands stressed by contracting muscle fibers, was able to account for the observations. A second model that provided the right predictions was a multiple impulse generator with cross-connections. Which of the two models best fits the experimental observations can be decided by comparing the calculated summation coefficients and cross-adaptation coefficients. A central impulse generator predicts a negative correlation, the multiple impulse generator a positive correlation. All of the observations were made using tendon organs of cat soleus muscle. Responses were recorded to stimulation of filaments of ventral root. In a comparison between 20 pairs of responses from six tendon organs the correlation between summation and cross-adaptation coefficients was found to be significantly positive. We conclude that the tendon organ model that accurately predicts all of the experimental observations incorporates multiple generators.
Design and Fabrication of a High Voltage Lightning Impulse Generator
