It has been proposed that lengthening of active muscle at long lengths is nonuniformly distributed between sarcomeres, with a few being stretched beyond overlap and most hardly being stretched at all. A small fraction of the overstretched sarcomeres may fail to reinterdigitate on subsequent relaxation, leading to progressive changes in the muscle's mechanical properties. Sartorius muscles of the toad Bufo marinus were subjected to repeated lengthening (eccentric) contractions at long lengths, while controls were passively stretched and then contracted isometrically or stretched at short lengths. The muscles undergoing eccentric contractions showed a progressive shift to the right of the length-tension curve, a fall in the yield point during stretch, an increase in slope of the tension response during stretch, and a fall in isometric tension. In control muscles, changes, if any, were significantly less. In electron micrographs, muscle fibers that had been subjected to a series of eccentric contractions showed sarcomeres with A bands displaced toward one half-sarcomere, leaving no overlap in the other half. Adjacent regions often looked normal. These results are all in agreement with the predictions of the nonuniform stretch of sarcomeres hypothesis.

1. Experiments were carried out to test the effect of changes in spindle resting discharge on the size of monosynaptic reflexes in the cat and on the H reflex in humans. Resting discharge was altered by contracting the triceps surae muscle at longer (hold-long) or shorter (holdshort) lengths than that at which the reflex was tested. 2. The reflex in the cat was larger after hold-long than after hold-short conditioning, and the difference, after an initial decline, was well maintained. For the human H reflex a similar pattern was observed except that 15 s after muscle conditioning the difference in reflex size had disappeared. 3. Monosynaptic reflex depression immediately after hold-long conditioning, when most of the muscle spindles are silent, was attributed to the high level of spindle discharge during the immediately preceding hold-long period. The time course of this inhibition was too long to be accounted for by presynaptic inhibition. 4. In the cat heteronymous muscle conditioning was used to test whether presynaptic inhibition could be responsible for reflex depression using the synergist muscle pair lateral gastrocnemius-soleus and medial gastrocnemius. Conditioning one of the pair did not affect the reflex in the other, the opposite result to that expected with presynaptic inhibition. A similar experiment in which the triceps H reflex in human subjects was facilitated by a quadriceps volley gave the same result. 5. Thus this study presents evidence that monosynaptic reflexes are depressed by the on-going discharge of muscle spindles in the homonymous muscle, but that this depression does not appear to involve 'classical' presynaptic inhibition.

has established that the responses of muscle spindles and their spinal reflex action are dependent on the muscle's immediate previous history of contraction and length changes, because muscle exhibits thixotropy (for a review see Proske et al. 1993). To demonstrate extremes of behaviour of spindles and spindle reflexes in relation to muscle history effects, we have devised two forms of conditioning, termed hold-long conditioning and hold-short conditioning. Hold-long conditioning consists of stretching the muscle to a length longer than that at which the test measurements are to be made, then contracting the muscle and holding it at that length for a few seconds before returning it to the test length. Similarly, hold-short conditioning is where the muscle is shortened, contracted and then stretched back to the test length. The reasoning underlying these forms of conditioning is as follows. As the muscle relaxes following a contraction, a number of stable cross-bridges form between actin and myosin filaments in sarcomeres of muscle fibres (Hill, 1968). The presence of stable cross-bridges gives the muscle a degree of passive stiffness. As a result, when the stiffened muscle fibres are shortened, as occurs after hold-long conditioning, they fall slack. By contrast, slack does not develop if the contraction is carried out at a shorter length and the muscle is then stretched back to the test length, that is, after hold-short conditioning. We hypothesize that slack is able to form in both extrafusal and intrafusal fibres. The presence of slack in intrafusal fibres has a large effect on the responses of muscle spindles.

Tendon jerk and H-reflexes are both potentiated by the Jendrassik manoeuvre, but the mechanism of potentiation remains uncertain. We investigated several possibilities in human subjects. Evidence for fusimotor activation during the Jendrassik manoeuvre was sought by recording the tendon jerk reflex as surface EMG in triceps surae after the muscles had been conditioned to leave their spindles in a slack, insensitive state. Interposing a Jendrassik manoeuvre between conditioning and the test reflex should have increased reflex amplitude by restoring spindle sensitivity, but this was not the case. In humans, a close synergist of the triceps surae is the quadriceps. A possible presynaptic disinhibitory mechanism was investigated by testing the effect of a Jendrassik manoeuvre on facilitation of the soleus H-reflex produced by a quadriceps afferent volley. The Jendrassik manoeuvre failed to increase facilitation, contrary to what would be expected if it reduced the level of tonic presynaptic inhibition; the assumption being that the inhibition acts on both homonymous and synergist afferent terminals. The Jendrassik manoeuvre did not increase the level of ongoing EMG in the soleus during a weak voluntary contraction, indicating that it does not operate by direct facilitation of motoneurones. There was found to be less potentiation of soleus tendon jerk and H-reflexes by the Jendrassik manoeuvre under conditions when spindles in the soleus were likely to have a high resting discharge rate. A remaining possibility is discussed: that the Jendrassik manoeuvre operates by modulation of oligosynaptic pathways that may contribute to the largely monosynaptic reflex response. These experiments demonstrate, with new, more sensitive methods than previously used, that neither is the fusimotor system involved in reinforcement nor are direct excitatory or presynaptic disinhibitory effects on motoneurones. While this confirms the previously prevailing view, none of the lingering uncertainties associated with the methods used now remains.

This is a study of the tendon jerk reflex elicited by a brief stretch applied to the triceps surae muscle group in the chloralose-anaesthetised cat. The size of the recorded reflex depended on stretch parameters (optimum at 300 microns amplitude at a rate of 100 mm/s) and on how the muscle had been conditioned. A reflex elicited after a conditioning contraction at the test length was often twice as large as after a contraction carried out at a length longer than the test length. This difference was attributed to the amount of slack introduced in the intrafusal fibres of muscle spindles by conditioning. The question was posed, did ongoing fusimotor activity exert any influence on the size of the tendon jerk? Depolarization indices (DPI) were calculated from responses of muscle spindles to stretch and correlated with the level of reflex tension. Values of DPI obtained from afferent responses with and without repetitive stimulation of identified fusimotor fibres suggested that with the stretch parameters used here the main influence of fusimotor activity was that it removed any pre-existing slack in muscle spindles and thereby increased reflex tension. In the absence of intrafusal slack, stimulation of static and dynamic fusimotor fibres had little additional influence on the size of the reflex. It is concluded that much of the variability typically seen with tendon jerks is due to muscle history effects. Since in muscles which have not been deliberately conditioned there is commonly some slack present in spindles, activity in fusimotor fibres is likely to reduce slack and therefore increase reflex size.