Reflexes Mediated by Non-Impulsive Afferent Neurones of Thoracic-Coxal Muscle Receptor Organs in the Crab, Carcinus Maenas

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Abstract
Address for reprints. A preparation of the thoracic-coxal muscle receptor organ of the posterior leg of the shore crab, in which central synaptic efficacy of the sensori-motor reflex pathways is maintained for long periods, is described. The reflex response to receptor muscle stretch commonly involves three promotor motoneurones, designated Pm1-3 in order of their recruitment. Motoneurone Pm1, and less frequently Pm2 and Pm3, may be tonically active during maintained receptor length changes within the in situ length range of the receptor muscle. The following observations suggest that the T rather than the S sensory fibre provides the afferent drive onto reflexly activated promotor motoneurones: selective section of the S or T sensory fibres; frequency ‘envelopes’ of individual motoneurone responses to trapezoid stretch stimuli, including features such as adaptation and velocity sensitivity of the reflex response; and the ‘hysteresis’ in the response to increasing followed by decreasing receptor length changes, with or without superimposed trapezoid stretch stimuli. The initial reflex response to ramp stretch can be directly related to the complex ‘initial component’ of the T fibre receptor potential waveform. This comprises a variable spiky alpha (α) component, followed by a longer duration, more predictable beta (β) component, which depends upon stimulus parameters such as stretch velocity and the length and tension of the receptor muscle at the onset of stretch. In the de-efferented receptor muscle, changes in compliance or ‘tonus’ resulting from receptor manipulation have a marked effect on the sensory, and hence reflex, response to stretch. As this would have profound implications for the functioning of this muscle receptor organ in vivo, a role for the receptor motor innervation in counteracting any such response variability seems likely.