Muscle Spindle - Sensitivity Modification

Sensitivity Modification

The function of the gamma motoneurons is not to supplement the force of muscle contraction provided by the extrafusal fibers, but to modify the sensitivity of the muscle spindle sensory afferents to stretch. Upon release of acetylcholine by the active gamma motoneuron, the end portions of the intrafusal muscle fibers contract, thus elongating the non-contractile central portions (see "fusimotor action" schematic below). This opens stretch-sensitive ion channels of the sensory endings, leading to an influx of sodium ions. This raises the resting potential of the endings, thereby increasing the probability of action potential firing, thus increasing the stretch-sensitivity of the muscle spindle afferents. For an interactive animation created by Jan Kowalczewski at the University of Alberta, demonstrating spindle afferent responses to muscle stretch with and without gamma (fusimotor) action, go to: Arthur Prochazka's Lab, University of Alberta

How does the central nervous system control gamma fusimotor neurons? It has been difficult to record from gamma motoneurons during normal movement because they have very small axons. Several theories have been proposed, based on recordings from spindle afferents.

• 1) Alpha-gamma coactivation. Here it is posited that gamma motoneurons are activated in parallel with alpha motoneurons to maintain the firing of spindle afferents when the extrafusal muscles shorten.
• 2) Fusimotor set: gamma motoneurons are activated according to the novelty or difficulty of a task. Whereas static gamma motoneurons are continuously active during routine movements such as locomotion, dynamic gamma motoneoruns tend to be activated more during difficult tasks, increasing Ia stretch-sensitivity.
• 3) Fusimotor template of intended movement. Static gamma activity is a "temporal template" of the expected shortening and lengthening of the receptor-bearing muscle. Dynamic gamma activity turns on and off abruptly, sensitizing spindle afferents to the onset of muscle lengthening and departures from the intended movement trajectory.