Regulation of muscle stiffness during periodic length changes in the isolated abdomen of the hermit crab.

Reflex activation of the ventral superficial muscles (VSM) in the abdomen of the hermit crab, Pagurus pollicarus, was studied using sinusoidal and stochastic longitudinal vibration of the muscle while recording the length and force of the muscle and the spike times of three exciter motoneurons. In the absence of vibration, the interspike interval histograms of the two larger motoneurons were bimodal; cutting sensory nerves containing most of the mechanoreceptor input removed the short interval peak in the histogram, indicating that the receptors are important in maintaining tonic firing. Vibration of the muscle evoked a reflex increase in motoneuron frequency that habituated after an initial peak but remained above control levels for the duration of stimulation. Motoneuron frequency increased with root mean square (rms) stimulus amplitude. Average stiffness during stimulation was about two times the stiffness of passive muscle. The reflex did not alter muscle dynamics. Estimated transfer functions were calculated from the fast Fourier transform of length and force signals. Coherence was >0.9 for the frequency range of 3-35 Hz. Stiffness magnitude gradually increased over this range in both reflex activated and passive muscle; phase was between 10 and 20 degrees. Reflex stiffness decreased with increasing stimulus amplitudes, but at larger amplitudes, this decrease was much less pronounced; in this range stiffness was regulated by the reflex. The sinusoidal frequency at which reflex bursts were elicited was approximately 6 Hz, consistent with previous measurements using ramp stretch. During reflex excitation, there was an increase in amplitude of the short interval peak in the interspike interval histogram; this was reduced when the majority of afferent pathways was removed. A phase histogram of motoneuron firing during sinusoidal vibration had a peak at approximately 110 ms, also suggesting that an important component of the reflex is via direct projections from the mechanoreceptors. These results are consistent with the hypothesis that a robust feedforward regulation of abdominal stiffness during continuous disturbances is achieved by mechanoreceptors signalling the absolute value of changing forces; habituation of the reflex, its high-threshold for low frequency disturbances and the activation kinetics of the muscle further modify reflex dynamics.