The responses of frog muscle spindles and fast and slow muscle fibres to a variety of mechanical inputs

1. The tension in the iliofibularis muscle of frogs was recorded while the muscle was stretched or released. At the same time recordings were made from single spindle afferents in dorsal root filaments. Either large or small motor nerve fibres were stimulated in split ventral root filaments.2. While small motor nerve fibres were stimulated the discharge from muscle spindle afferents was greatly increased by stretching, and greatly reduced by shortening the muscle. This sensitivity to movement was shown even if the movements were small, so that a stretch of 0.2% of the muscle length was sufficient to cause a pronounced increase in the afferent discharge.3. In contrast, during stimulation of the large motor nerve fibres the spindle was much less sensitive to movements with the result that even stretches or releases of the muscle by 1 mm did not cause very large changes in the discharge frequency.4. The tension in slow extrafusal muscle fibres in many ways mirrored the spindle discharge during the stimulation of small motor nerve fibres, for the tension was greatly increased by stretching, even through small distances, and greatly reduced by releasing the muscle. The tension in fast extrafusal muscle fibres was much less changed by such movements, and thus was rather like the spindle discharge during stimulation of large motor nerve fibres.5. As the extrafusal muscle fibres do not directly pull on and excite the spindle afferents, the simplest explanation for the similarities between the muscle tension and the spindle discharge is that the mechanical properties of the intrafusal muscle fibres innervated by the large motor nerve fibres are like those of fast extrafusal muscle fibres, and that the mechanical properties of the small intrafusal fibres are similar to those of slow extrafusal muscle fibres.6. It is shown that the cross-bridge sliding filament mechanism of muscle contraction provides a ready explanation for the differences found between fast and slow muscles, and it is concluded that a most important functional difference between the two sorts of intrafusal muscle fibres is the speed of their contractions, for it is this which determines their contrasting actions on the spindle.7. It was also found that low rates (< 4/sec) of small motor nerve fibre stimulation were often very effective in exciting the spindles. These rates produced rather little extrafusal tension.     

An electrophysiological analysis of responses from lizard muscle spindles

1. Stretch receptor discharges were recorded in the lizard Tiliqua nigrolutea. Responses were identified as coming from muscle spindles by their ;in parallel' behaviour during a twitch.2. Recordings from spindles in the iliofibularis muscle which contains many single innervated fibres revealed spindles which discharged on the rising phase of the muscle contraction. This ;in series' response increased in frequency and duration at high resting tensions on the muscle and was attributed to a specific intrafusal contraction.3. The ;in series' discharge could be induced by stimulation of only one small filament of the muscle nerve. This suggested a focal intrafusal motor innervation.4. Responses from spindles in the semitendinosus muscle, which contains many multiple innervated fibres provided examples of an ;in series' response which could be produced by stimulation of several muscle nerve filaments. This was interpreted as being an example of multiple intrafusal motor innervation.5. Spindles with short or elongated sensory regions were subjected to ;ramp' stretches. The presence of distinct phasic and tonic responses from the two types was confirmed and related to the behaviour of mammalian spindles.     

A comparison of the spindles in two different muscles of the frog

1. The responses of spindles in the iliofibularis muscle of frogs to stretch during either small motor nerve fibre stimulation or the application of suxamethonium were compared.2. All spindles which were excited by small motor nerve fibre stimulation were also excited by suxamethonium, and their responses to these two methods of excitation were very similar. The drug dose was usually 5-10 mug/ml. but smaller and larger doses were effective. Large doses (> 100 mug/ml.) could sometimes lead to a reversible partial block of the spindle response to stretch.3. Suxamethonium also caused a prolonged contraction in extrafusal slow muscle fibres. This contraction was not responsible for the effect on the spindle, because the time course of its action on the muscle tension and on the spindle afferent was different.4. It was concluded that suxamethonium stimulated prolonged contraction in the small intrafusal muscle fibres, which are known to be innervated by the small motor nerve fibres.5. Only about half of the spindles in the iliofibularis muscle were excited by suxamethonium.6. In the sartorius muscle which has no slow extrafusal muscle fibres, no spindles were found to be excited by suxamethonium in the way characteristic of that due to small intrafusal muscle fibre contraction.7. It is concluded that, in frog muscles which have no slow extrafusal fibres, the muscle spindles do not have small intrafusal muscle fibres of the kind found in the iliofibularis muscle.     

Aftereffects in the responses of cat muscle spindles

Responses have been recorded from primary endings of muscle spindles in the cat soleus muscle. Changes in spindle responsiveness were measured following a period of conditioning that consisted of a series of rapid stretches or of tetanic ventral root stimulation. In the testing procedure the response of a single spindle afferent was recorded to stimulation of a dynamic fusimotor axon during a slow stretch. Changes in gross afferent discharge coming from the muscle were measured by integrating the activity recorded in dorsal roots. If, after conditioning stretches, the muscle was immediately returned to its initial length, the spindle responded to the test fusimotor stimulation with a high-frequency burst of afferent impulses. If the muscle was held stretched for 3 s after conditioning the response to the brief test tetanus was small or "depressed." It has been suggested that conditioning stretches result in detachment of stable crossbridges in intrafusal fibers and that these bridges then reform over the next few seconds at whatever length the muscle happens to have at the time. When it is long, shortening the muscle back to the initial length leads to the development of slack in intrafusal fibers because of the passive stiffness they have acquired from the presence of the stable bridges. Under these conditions a brief test fusimotor tetanus will lead to a depressed response because the slack must first be taken up before a full response can be generated. It was possible to reverse the depression by interposing an extrafusal contraction during the period between the conditioning and test sequences. It is suggested that lateral compression from the contracting extrafusal fibers and the stretch they impose as they relax reduces any intrafusal slack and thereby reduces the depression. A more quantitative measure of intrafusal slack than the test for depression is to determine the delay in onset of the afferent response to a longer fusimotor tetanus. The delay was short a long initial muscle lengths where, if the muscle was left undisturbed, it soon disappeared completely and spontaneously. It is suggested that at long lengths passive tension in the muscle tends to remove any slack in intrafusal fibers and therefore removes any after effects. The rise in resting discharge of muscle afferents after a conditioning tetanus applied to the ventral root ("postcontraction sensory discharge") can be accounted for by the same hypothesis.(ABSTRACT TRUNCATED AT 400 WORDS)     

The responses of frog muscle spindles during stimulation of slow motor axons

Summary Responses of muscle spindles of the iliofibularis muscle of the frogLitoria aurea have been recorded during single shock and repetitive stimulation of single functional motor axons. Repetitive stimulation of axons which innervated slow muscle, and on four occasions, axons which innervated twitch muscle, produced a large increase in the dynamic response of the spindle to a ramp-and-hold stretch. While extrafusal slow muscle did not respond to a single motor volley, some spindles did, especially if at the same time the muscle was being stretched. In an explanation of the effect of muscle stretch on responses of spindles to slow motor volleys it was proposed that stretch acted to reduce the internal motion in muscle fibres produced by a non-uniform distribution of sarcomere lengths. It was proposed that this kind of effect may account for dynamic fusimotor actions in all vertebrate spindles.Financial assistance was provided by a grant from the National Health and Medical Research Council of Australia, Grant No. 78/5721     

Responses of muscle spindles following a series of eccentric contractions

To investigate the effects of eccentric exercise on the signalling properties of muscle spindles, experiments were done using the medial gastrocnemius muscle of cats anaesthetised with 40 mg/kg sodium pentobarbitone, i.p. Responses were recorded from single afferent nerve fibres in filaments of dorsal root during slow stretch of the passive muscle and during intrafusal contractions at a range of lengths, before and after a series of eccentric contractions. The sensitivity to slow stretch was measured as the average firing rate between muscle lengths 10.5 and 9.5 mm shorter than the physiological maximum (L_m), during stretch at 1 mm/s over the whole physiological range. The mean sensitivity of both primary and secondary spindle endings increased slightly, but not significantly, after a series of 20–150 eccentric contractions consisting of a 6 mm stretch, at 50 mm/s, to a final length of between L_m –7 mm and L_m, during stimulation of the whole muscle or sometimes of single fusimotor fibres. Discharges were recorded from primary endings during fusimotor stimulation at 100–150 pulses/s, and from secondary endings during static bag intrafusal contractures produced by i.v. injection of 0.2 mg/kg succinyl choline. Spindle responses were recorded, over a range of muscle lengths, in steps covering the whole physiological range. About half of the responses showed a peak in the relation between length and net increase in firing rate, while the remainder either progressively increased or progressively decreased over the physiological range. No large or consistent changes were seen after the eccentric contractions. It is concluded that the intrafusal fibres of muscle spindles are not prone to damage of the kind seen in extrafusal fibres after a series of eccentric contractions.     

Innervation of extrafusal and intrafusal fibres in snake muscle

1. Intrafusal fibres of snake receive motor supply from branches of axons innervating extrafusal motor units. By intramuscular stimulation of motor units by single shocks, and critical curarization of the muscle, we have identified at least some of the motor units contributing motor supply to individual intrafusal fibres. Intrafusal fibre activation was observed by visual examination of the contracting intrafusal fibre, and by recording the resulting spindle afferent discharge.2. The main finding is that in some cases the motor supply to one intrafusal fibre comes from more than one motor unit. The contributing motor units may be either dissimilar twitch units, or twitch and tonic units. Thus some of the intrafusal fibres studied showed polyneuronal motor innervation of heterogeneous origin.3. In critically curarized muscle, the time course of a spindle afferent discharge, following single-shock stimulation of a motor unit contributing motor supply to the intrafusal fibre, showed little variation with the type of motor unit being stimulated.4. The response of each spindle to a standard stretch was recorded. There was no correlation between dynamic index and type of motor unit or units contributing motor supply. However, the method limits the value of negative findings, and this is discussed.5. The contraction times and tensions of a sample of motor unit isometric twitches are described.     

A muscle spindle model for primary afferent firing based on a simulation of intrafusal mechanical events.

1. A muscle spindle model for primary afferent firing is presented that contains two components representing a gamma d-dependent (bag1) and gamma s-dependent (bag2/nuclear chain) intrafusal fiber. Each of the intrafusal fibers is composed of a linear elastic element representing the sensory part and a muscle fiber representing the muscular part. 2. The muscular part of the bag1 was modeled as a slow twitch, that of the bag2 as a fast twitch muscle fiber. 3. The sensory regions were linear length transducers, generating a rising depolarization on increasing stretch. The input of both bags was fused by taking the largest depolarization to determine a generator potential. The rate of primary afferent firing depended on this generator potential as well as on its rate of change. 4. To simulate the high sensitivity of muscle spindles to small amplitudes of stretching, a model analogue of cross-bridge fixation (or stiction) has been included in the muscular part of the bag1 fiber. This makes use of one hundred cross-bridge regions that release one after the other, provided a certain breaking force is exceeded. 5. The values of the mechanical parameters that defined the model were selected by a computerized search procedure. 6. The values found by means of this procedure allowed the model to provide an accurate simulation of experimental data on ramp-and-hold stretches (for 6 different stretch velocities under variable conditions of fusimotor activity). 7. On sinusoidal stretches at a frequency of 1 Hz the spindle model responded with about one-half the discharge modulation reported in experimental studies. Its phase advance tended to be slightly lower than that observed for real spindles. 8. Frequency response curves showed the same high sensitivities at high frequencies as those observed in real spindles. 9. Close evaluation of the model compared with experimental results in literature reveal its merits as well as its limitations. Because the model is structural rather than phenomenologic, it provides insight into how intrafusal events may contribute to observed firing properties of real muscle spindles.     

Changes in size of the stretch reflex of cat and man attributed to aftereffects in muscle spindles

1. This is a report of experiments carried out on the cat and on man, which demonstrate that conditioning of a muscle by contraction and movement can lead to changes in amplitude of stretch reflexes elicited in that muscle. 2. In triceps surae of the cat, the reflex response to a brief stretch was recorded after conditioning with a whole-muscle contraction followed by a pause at a length either 5 mm longer or shorter than the length at which the reflex was elicited. Following conditioning at the long length the reflex response was less than half as large as that following conditioning at the short length. 3. The changes in reflex amplitude could be correlated with an altered stretch responsiveness of muscle spindles in the soleus muscle. When the muscle had been held long during conditioning, a subsequent brief stretch applied at an intermediate length elicited fewer impulses in primary endings of spindles than after conditioning at a short length. 4. The same kind of experiment was then carried out on adult human subjects. When a tendon tap was applied to the Achilles tendon after a voluntary contraction and relaxation of triceps surae with the muscle at a long length, (foot dorsiflexed) the reflex was frequently less than half the size it had been after a contraction at a short length (foot plantarflexed). It was concluded that the same kind of spindle aftereffects as observed for cat soleus spindles were responsible for the changes in reflex amplitude. 5. It was found both in the cat and in human subjects that the changes in reflex amplitude after conditioning became progressively less as the test length was made longer. 6. The explanation put forward to account for these observations is that stable cross-bridges form between actin and myosin filaments of passive intrafusal (and extrafusal) fibers. When the muscle is shortened several seconds after a contraction at a long length, the intrafusal fibers, stiffened by the presence of cross-bridges, fall slack. Slack does not develop after a contraction at a short muscle length, as the fiber is stretched to the test length. Since any slack must first be taken up by the test stretch, there is a smaller afferent response and consequently a smaller reflex contraction in response to a tendon tap after conditioning at a long length.(ABSTRACT TRUNCATED AT 400 WORDS)     

The responses of secondary endings of cat soleus muscle spindles to succinyl choline

This report describes the effects of succinylcholine (SCh) on the secondary endings of cat soleus muscle spindles and attempts to explain them in terms of the action of the drug on intrafusal fibres. All but 2 of 41 secondary endings studied in detail showed a significant response to a single intravenous injection of 200 g kg^-1 SCh. This consisted of a rise in the resting rate or development of a resting discharge if the spindle had previously been silent and an increase in the response to stretch. The increases in the responses to stretch were weaker than those observed for primary endings of spindles, but were much larger than those of tendon organs, which showed very little effect with this concentration of drug. The response to SCh showed two features consistent with its action being mediated via an intrafusal muscle fibre contraction rather than a direct depolarising action on the afferent nerve ending. In the presence of SCh, secondary endings were able to maintain a discharge during muscle shortening at rates, on average, more than 5 times greater than under control conditions. Secondly, the increase in spindle discharge produced by SCh showed a length dependence similar to that for fusimotor stimulation. Further support for the action of SCh being principally via an intrafusal fibre contraction was provided by the observation that its effects were abolished by the neuromuscular blocker gallamine triethiodide. The time course of recovery of SCh responses, following their blockade by gallamine, was much slower than recovery of extrafusal tension and closely paralleled that for the recovery of fusimotor responses. In three separate experiments on the medial gastrocnemius muscle the possibility that SCh may exert an excitatory action on spindle sensory endings through the liberation of potassium ions from the muscle was tested by tetanic stimulation of the muscle. This had no detectable excitatory effect. Several observations were made on the effect of SCh on responses of cutaneous receptors. SCh did not change levels of spontaneous activity or responses to mechanical stimulation of either slowly or rapidly adapting mechanoreceptors. It was argued for both tendon organs and cutaneous receptors that if SCh had a direct action on the nerve ending at the concentrations used here, some responses of these receptors to the drug might have been expected. All of the above supports the view that secondary endings of spindles are able to respond to SCh by the development of an intrafusal fibre contracture. The question of the intrafusal fibre types involved is discussed.     

After-effects of stretch on the responses of cat soleus muscle spindles to static fusimotor stimulation

Summary Mammalian muscle spindles show persistent after-effects following conditioning stretch or fusimotor stimulation. Most previous observations have been carried out on primary endings of spindles and using dynamic fusimotor stimulation. We report here observations on after-effects produced either by conditioning stretch or by static fusimotor stimulation on the responses of primary and secondary endings to a slow test stretch during which a brief burst of static fusimotor stimulation is applied. We find that the response to the test burst is large if the muscle is kept short after conditioning but it becomes depressed if the muscle is held stretched for 3 s following conditioning. We attribute these effects to the presence of stable cross-bridges between actin and myosin filaments in intrafusal fibres. We conclude that, qualitatively, after-effects using static fusimotor testing are the same as with dynamic fusimotor testing and this must be taken into account when providing an explanation for the phenomenon.     

After-effects of fusimotor stimulation on the response of muscle spindle primary afferent endings

1. The experiments were performed on the soleus muscle of the anaesthetized cat in which the ventral roots had been cut.2. A short period of repetitive stimulation of a single fusimotor fibre which influenced a particular spindle primary ending invariably caused a characteristic alteration in the response of the same ending to a subsequently applied ramp stretch of the muscle. The change consisted in the appearance of a burst of impulses at the beginning of the stretch where none had been present before. Occasionally, such an ;initial burst' was spontaneously present; it was then enhanced following fusimotor stimulation.3. This after-effect of fusimotor stimulation was abolished by a subsequent stretch of the muscle, but otherwise persisted for over a minute.4. When the muscle was released to below the length at which the spindle had been facilitated and a testing stretch applied from the new initial length there was no burst of impulses at the beginning of stretch. There was, however, a burst as the muscle was stretched through the length at which the fusimotor fibre had been stimulated.5. These effects are suggested to be due to the persistence of stable bonds between the actin and myosin filaments of the intrafusal fibres, so that their previously activated regions were ;stuck' at the length they were when the fusimotor stimulation was applied.6. Such effects were produced both by static and by dynamic fusimotor fibres. The effects of the two kinds of fusimotor fibre, however, appeared to be mediated by different intrafusal muscle fibres. This was shown by stimulating one kind of fibre with the muscle slightly stretched, then releasing the muscle a few mm and stimulating the other kind of fibre to the same spindle. A subsequent testing stretch then elicited two bursts, one at the beginning and one in the middle of the stretch.     

Muscle spindle discharge in response to contraction of single motor units

1. In human subjects, microelectrode recordings were made from 25 muscle spindle afferents and two tendon organ afferents coming from muscles innervated by the peroneal nerve. 2. Stimulation at low intensity through the recording microelectrode activated efferent axons innervating motor units in close proximity to the muscle spindle or tendon organ. There was a clear alteration in the discharge of 17 afferents (15 muscle spindle, 2 tendon organ) in response to twitch contractions that involved only one, two, or three motor units. With three other afferents there was a less overt but statistically significant alteration in discharge rate by the twitch contraction of a single motor unit. 3. The sensitivity of 21 receptors (20 spindles, 1 tendon organ) to twitch contractions of anatomically close motor units was contrasted with their sensitivity to twitches of more remote motor units in the muscle. In no instance was the sensitivity to the contraction of remote motor units greater than that to the contraction of local motor units stimulated through the microelectrode; with remote stimulation many units usually had to be activated before the resulting twitch contraction altered the discharge of an afferent. 4. It is concluded that muscle spindles as well as tendon organs can play a role in monitoring the activity of motor units anatomically close to the receptor.     

Two kinds of resting discharge in cat muscle spindles.

1. The behavior of primary endings of cat soleus muscle spindles was studied during shortening steps carried out at different muscle lengths. 2. Spindles were of two kinds: one, silent spindles, whose afferents fell silent after the shortening, at least over part of the range of lengths tested. The second, spontaneous spindles, resumed firing at all lengths. 3. For silent spindles, the duration of the silent period, measured at lengths where they did recover a resting rate, depended directly on muscle length and became shorter at longer lengths. This is what would be expected if the slack introduced in the spindle by the shortening step was removed more rapidly at longer lengths by the higher passive tension. For spontaneous spindles, on the other hand, the duration of the silent period after the shortening was largely independent of muscle length and depended on the spindle's rate of firing immediately before the shortening. 4. At intermediate lengths the discharge of slack spontaneous spindles remained unaffected by an isometric muscle contraction. It was therefore not possible to produce a pause in the discharge, behavior normally taken as typical of spindles. The discharge could be interrupted by the contraction if this was combined with a large shortening movement. 5. It is proposed that when intrafusal fibers are slackened by a shortening step, the resting discharge in spontaneous spindles is generated by a maintained depolarization of the annulospiral ending resulting from extension of the terminal coils by forces from within the receptor. A shortening contraction compresses the spirals to interrupt the discharge. The sensory endings of silent spindles remain below threshold until the spirals have been opened out sufficiently by external stretch.     

A determination of static mechanical properties of intrafusal muscle in isolated cat muscle spindles.

The sensitivity of mammalian muscle spindles to stretch is greater in a stretched muscle than in a slack one. We have investigated this behavior in isolated muscle spindles removed from cat tenuissimus muscle. We measured the steady-state strain of intrafusal muscle in sensory and non-sensory regions and found that there is a proportional relationship between sensory strain and receptor sensitivity; both increase with spindle length. By comparing intrafusal strain of sensory and non-sensory areas with and without an intact spindle capsule, we conclude that capsule does not contribute to the non-linear sensitivity. Measurements of steady-state tension indicate that the striated portions of the intrafusal muscle exhibit a non-linear stiffness which can quantitatively account for the observed behavior.     

Small-signal analysis of response of mammalian muscle spindles with fusimotor stimulation and a comparison with large-signal responses.

1. Response dynamics of primary and secondary muscle spindle endings to small-amplitude sinusoidal stretches were found to be unaltered by tonic repetitive stimulation of fusistatic or fusidynamic fibers. 2. Overall sensitivity of these receptors is decreased by fusistatic stimulation and either unchanged, increased, or decreased by fusidynamic stimulation at rates of 75/s or greater. 3. In the case of primary endings, the results obtained with small-amplitude sinusoidal stretches are not compatible with the response of these receptors to large-amplitude ramp stretches. The difference is explained by dependence of receptor dynamics on stretch amplitude. Fusistatic stimulation tends to prevent those changes in dynamics, whereas fusidynamic stimulation tends to enhance them. 4. In the case of secondary endings, the results obtained with small- and large-amplitude stretches appear to be compatible with a linear model for this receptor (i.e., one with dynamics independent of input parameters). 5. By modulating the frequency of stimulation applied to fusimotor fibers and comparing the resulting afferent response to the receptor response to stretch dynamic characteristics of intrafusal muscle contraction can be deduced. The results suggest that the dynamics of fusiastatic and fusidynamic contraction are the same and, furthermore, that they are the same as those of extrafusal muscle. We note that the result is incompatible with measurements of the time course of twitch and tetanus development and suggest, therefore, that muscle dynamics are a function of contractile state.     

Structure, distribution and innervation of muscle spindles in avian fast and slow skeletal muscle

Muscle spindles in 2 synergistic avian skeletal muscles, the anterior (ALD) and posterior (PLD) latissimus dorsi, were studied by light and electron microscopy to determine whether morphological or quantitative differences existed between these sensory receptors. Differences were found in the density, distribution and location of muscle spindles in the 2 muscles. They also differed with respect to the morphology of their capsules and intracapsular components. The slow ALD possessed muscle spindles which were evenly distributed throughout the muscle, whereas in the fast PLD they were mainly concentrated around the single nerve entry point into the muscle. The muscle spindle index (number of spindles per gram wet muscle weight) in the ALD was more than double that of its fast-twitch PLD counterpart (130.5±2.0 vs 55.4±2.0 respectively, n=6). The number of intrafusal fibres per spindle ranged from 1 to 8 in the ALD and 2 to 9 in the PLD, and their diameters varied from 5.0 to 16.0 μm and 4.5 to 18.5 μm, respectively. Large diameter intrafusal fibres were more frequently encountered in spindles of the PLD. Unique to the ALD was the presence of monofibre muscle spindles (12.7% of total spindles observed in ALD) which contained a solitary intrafusal fibre. In muscle spindles of both the ALD and PLD, sensory nerve endings terminated in a spiral fashion on the intrafusal fibres in their equatorial regions. Motor innervation was restricted to either juxtaequatorial or polar regions of the intrafusal fibres. Outer capsule components were extensive in polar and juxtaequatorial regions of ALD spindles, whereas inner capsule cells of PLD spindles were more numerous in juxtaequatorial and equatorial regions. Overall, muscle spindles of the PLD exhibited greater complexity with respect to the number of intrafusal fibres per spindle, range of intrafusal fibre diameters and development of their inner capsules. It is postulated that the differences in muscle spindle density and structure observed in this study reflect the function of the muscles in which they reside.     

Slowing of the discharge of secondary endings of cat muscle spindles during fusimotor stimulation

Summary Responses of secondary endings of muscle spindles of the peroneus tertius muscle of the anaesthetized cat have been recorded during repetitive stimulation of functionally single fusimotor fibres that produced slowing of the discharge. In a sample of 125 pairs of single fusimotor fibres and secondary spindle afferents 5 examples of slowing were seen. The amount of slowing became less at longer muscle lengths. Conditioning the spindle by stimulating the muscle nerve at fusimotor strength, at a length 2.5 mm longer than the test length, and then returning to the test length 3 seconds later led to a greater degree of slowing of the discharge than after conditioning stimulation at the test length. With one exception, responses to muscle stretch were reduced during stimulation of a fusimotor fibre that produced slowing. On two occasions stimulating a fusimotor fibre that produced slowing of the response of one secondary ending, led to excitation of two other endings. Two possible explanations for the generation of slowing responses have been considered. The first is that the slowing is the result of contraction of the region of intrafusal fibre directly underlying the secondary sensory ending. The second, which we favour since it accounts for the facts more adequately, is that slowing is the result of shortening of the region of nuclear chain fibres on which the sensory ending lies, produced by movement in an adjacent nuclear bag fibre.     

Sympathetic innervation of human muscle spindles

The aim of the present study was to investigate the presence of sympathetic innervation in human muscle spindles, using antibodies against neuropeptide Y (NPY), NPY receptors and tyrosine hydroxylase (TH). A total of 232 muscle spindles were immunohistochemically examined. NPY and NPY receptors were found on the intrafusal fibers, on the blood vessels supplying muscle spindles and on free nerve endings in the periaxial space. TH‐immunoreactivity was present mainly in the spindle nerve and vessel. This is, to our knowledge, the first morphological study concerning the sympathetic innervation of the human muscle spindles. The results provide anatomical evidence for direct sympathetic innervation of the intrafusal fibers and show that sympathetic innervation is not restricted to the blood vessels supplying spindles. Knowledge about direct sympathetic innervation of the muscle spindle might expand our understanding of motor and proprioceptive dysfunction under stress conditions, for example, chronic muscle pain syndromes.     

Afferent fibres from muscle receptors in the posterior nerve of the cat's knee joint

Summary The properties of some receptors with afferent fibres in the cat's posterior knee joint nerve have been examined, especially those discharging tonically with the joint in intermediate positions between full flexion and extension. Some of these receptors behave like muscle spindles, and respond to manoeuvres which stretch popliteus muscle. Both in single unit and whole nerve recordings their discharge pauses during a popliteus twitch, and can be strikingly augmented by tetanic stimulation of a number of popliteus fusimotor fibres isolated from ventral root filaments. The action of succinylcholine on these receptors closely resembles its effect on popliteus spindle units with fibres sited normally in the popliteus nerve. Other units with properties suggesting origin from popliteus tendon organs were also observed; their fibres and those of the spindle units conducted at Group I velocity. It is concluded that some afferent fibres from popliteus spindles and possibly tendon organs commonly pursue an aberrant course in the posterior articular nerve of the knee joint.