Dynamics of reflex cocontraction in hermit crab abdomen: experiments and a systems model

Abstract

1. Both stretch and release of the ventral superficial muscles (VSM) in the abdomen of the hermit crab, Pagurus pollicarus, activate the VSM motoneurons in the intact animal and in the isolated abdomen. 2. This reflex was studied by recording intracellularly from muscle fibers innervated by single motoneurons during stretch and release of the VSM. The three motoneurons of the right fourth segment respond to both stretch and release with a phasic burst lasting approximately 250 ms. The burst in the two tonic motoneurons has two components, a short burst lasting 10-20 ms, with a latency from the beginning of stretch of 60-90 ms, and a longer burst of variable length, with a latency of 120 ms. Ramp stretches of different amplitudes and velocities were used to show that the first component is proportional to the absolute value of the second derivative of force and the second component to the absolute value of the first derivative of force. 3. Stretch and release of the VSM also simultaneously evoke phasic bursts in the motoneurons of the dorsal superficial muscles and the VSM circular muscles (functional antagonists of the longitudinal VSM), as well as in contralateral homologues of the same segment and in ipsilateral homologues of the next anterior segment. The effect of this coactivation is to stiffen the abdomen in response to perturbations in any direction. 4. Stretch or release of phasic mechanoreceptors in the VSM evokes this reflex. Isometric electrical stimulation of the isolated muscle also activates them, showing that they are transducing changes in force and suggesting that they operate to increase muscle stiffness by positive feedback. 5. A mathematical systems model of this reflex, composed of two parallel pathways activating the motoneurons, was constructed. The first pathway produces a signal proportional to the absolute value of the second derivative of force, the second pathway a signal proportional to the first derivative of force. The sum of the signals from the two pathways is filtered by an adaptation process, which is followed by a low-pass filter representing muscle activation kinetics. The muscle activation signal is then fed back to multiple muscle force. 6. Simulations using this model generate the phasic bursts to stretch and release as well as reproducing the frequency dependence of this reflex. The predominant action of this reflex is to enhance muscle stiffness.