Neural signals from proprioceptors in muscles provide length and force-related linkages among muscles of the limbs. The functions of this network of heterogenic reflexes remain unclear. New data are reported here on the distribution and magnitudes of neural feedback among quadriceps and triceps surae muscles in the decerebrate cat. The purpose of this paper was to distinguish whether inhibitory-force feedback is directed against muscles by virtue of the motor-unit composition or articulation of the muscle. These studies were carried out using controlled stretches and measurements of the resulting force responses of individual quadriceps and triceps surae muscles. Responses were evoked over a wide range of background force levels. In agreement with earlier electrophysiological studies, excitatory length feedback strongly linked the vastus muscles, but excitatory reflexes between each vastus and rectus femoris muscles were weak. We also observed a substantial excitatory linkage from the vastus muscles to the soleus muscle. In contrast, force-related inhibition was absent in the heterogenic reflexes among the vastus muscles but strong and bidirectional between each vastus muscle and the rectus femoris muscle and between triceps surae and quadriceps muscles. We conclude that short-latency feedback in the hindlimb is organized according to muscle articulation. Length feedback within muscle groups regulates joint stiffness while interjoint length feedback may compensate for the effects of nonuniform inertial properties of the limb. Force feedback is organized to regulate coupling between joints and, along with length feedback, determine the mechanical properties of the endpoint.
Healthy subjects were asked to make a voluntary ramp and hold contraction. The duration of the ramp stage was 500 ms, and the torque increment in this period was set to 15 Nm. The contraction was made from a relaxed and from a 5 Nm background torque situation. Hoffmann (H-) reflexes were elicited during the voluntary contraction, mostly with 100 ms intervals. These experiments showed an increase (facilitation) in the H-reflex before the torque or the EMG started to increase. This facilitation of the H-reflex remained during all the stages of the voluntary movement and declined to normal levels again only at the very end of the hold phase, which lasted for one second. This specific pattern of facilitation during a voluntary contraction was modeled using a modeling language, that is specifically designed to calculate neuronal systems with a high degree of reality (Ekeberg et al., 1991). Our model consisted of a motoneuron pool with 200 neurons connected to an EMG-model of the human soleus muscle and an extra group of higher-level neurons for controlling the amount of decrease of presynaptic inhibition. The model was used to simulate the observed modulation of the H-reflex with both a presynaptic and a postsynaptic mechanism. Simulations showed that a continuous change in the descending control signals is needed to make the model based on postsynaptic mechanism fit with the experimental data, whereas no extra control from the CNS over the excitatory drive to the motoneuron pool is needed when the decrease of presynaptic inhibition mechanism is applied.
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