Responses Of Primary Endings Research Articles

Development of the sensory innervation of muscle spindles in the kitten

This is a report of the changes observed in the pattern of sensory innervation of muscle spindles in hindlimb muscles of kittens during the first four weeks of life. The structural analysis, made on teased, silver-stained preparations, was complemented by a series of recordings of afferent responses of kitten spindles during ramp-and-hold stretches of the muscle. The primary endings of spindles from newborn animals showed a large degree of variability in their branching pattern and branches formed a network across the intrafusal fibres. In older animals there was less variability and lateral branches of stem axons began to encircle the intrafusal fibres. The process of maturation was characterized by a more uniform shape of the endings and more complete, evenly spaced, annulospiral terminals. Recordings of the responses of primary endings of spindles during muscle stretch showed that several features of the adult response were already present in the newborn, although the overall rate of discharge was very much lower. It was concluded that the changes observed in the structure of the sensory endings of kitten spindles did not have clearly identifiable physiological correlates. It appears that an annulospiral shape of the sensory terminals is not a necessary prerequisite for the generation of stretch responses. The predominant factor which appears to determine the responses of spindles to stretch is the maturity of the intrafusal fibres, in particular, the bag2 fibre.

Responses of primary endings of cat muscle spindles to locally applied vibration

Responses of muscle spindles in the cat soleus muscle have been studied during vibration applied locally to the belly of the muscle. Bursts of vibration at 170 Hz and with a peak amplitude of 200 microns were applied to a site at which local pressure initiated impulses from the spindle. The response to vibration depended on the conditioning of the muscle immediately beforehand and the placement of the vibrator. The length at which the vibration was applied was called the test length; this was typically 10 mm less than the muscle's maximum length in the body. After a fusimotor strength contraction at a length 2-5 mm longer than the test length, vibration sensitivity, measured on return to the test length, was low. If the muscle was contracted at a length 2-5 mm shorter than the test length, vibration sensitivity was high. The low vibration sensitivity following conditioning at the longer length was attributed to the development of slack in intrafusal fibres. In the presence of slack, stimulation of some static fusimotor fibres was able to restore vibration sensitivity fully. It is suggested that the vibration sensitivity of passive spindles arises largely in bag2 intrafusal fibres.

The distribution of static gamma‐axons in the tenuissimus muscle of the cat.

1. The distribution of static gamma-axons within and among muscle spindles of the tenuissimus muscle has been studied in the anaesthetized cat, on the basis of the effects on the responses of primary endings when bag2 or chain fibres or both are activated by static gamma-stimulation. 2. Locations of spindles were marked for subsequent histological analysis using teased, silver-impregnated preparations. 3. Static effects were classified into: (i) biassing; (ii) driving; or (iii) indeterminate categories. 4. Critical correlations established that the biassing type was produced by bag2 activity, either alone or in combination with chain fibres, whereas the driving type was produced by chain fibres active alone. Indirect evidence suggested that indeterminate effects were produced by bag2 and chain fibres active together. 5. The static gamma-axons showed some differential distribution according to their conduction velocities: faster-conducting axons were likely to be more widely distributed among spindles but less likely to innervate chain fibres alone than were more slowly conducting axons. 6. The results are discussed in terms of their possible functional and developmental significance.

Activation of cat muscle spindles by static skeletofusimotor axons.

The discharges from primary and secondary spindle endings of the cat peroneus tertius muscle were recorded during stimulation of static skeletofusimotor (static beta) axons at frequencies comparable with the presumed range of motoneuronal firing rates. When stimulated at 20-40/s, static beta-axons exerted typical static actions on the spindles they innervated, including activation of primary endings with reduction of their dynamic sensitivity and activation of secondary endings. For these frequencies, the extrafusal portions of static beta-motor units developed unfused contractions producing oscillations of tension within the muscle. After suppression of extrafusal contractions, the effects of the stimulation of static beta-axons on spindle discharge could persist unaltered, showing that extrafusal events need not interfere with the specific intrafusal actions of static beta-axons. Stimulation of a static beta-axon at 20-40/s often elicited a response of primary endings in which the discharge exactly followed the stimulation frequency, i.e. it was driven 1:1. Purely mechanical excitation of a spindle by unfused extrafusal contractions could also drive its discharge at the stimulation frequency. The persistence of static beta-driving after suppression of extrafusal contraction provided evidence for its intrafusal origin. Driving elicited by static beta-axons could persist during changes in muscle length, but small fluctuations in the delay between each impulse and the preceding stimulus were observed. These fluctuations were clearly related to the changes in muscle length, indicating that although the primary ending discharge remained driven 1:1 at the stimulation frequency, the receptor was not totally insensitive to length changes.

Effects of slow muscle stretch on the responses of primary and secondary endings to small amplitude periodic stretches in de-efferented soleus muscle spindles

The responses of primary and secondary endings of de-efferented soleus spindles to small amplitude periodic stretches superimposed on slow ramp stretches have been studied. For primary endings, the responses become progressively larger especially during the last period of the ramp whereas for secondary endings, after an initial period of moderate growth, the amplitude of the responses display a relative reduction precisely when the responses of the primary endings are the largest. These differences can be interpreted in terms of progressive increase in stiffness of the striated polar portions of the intrafusal muscle fibers.