Comparison of the effects of stimulating groups of static gamma axons with different conduction velocity ranges on cat spindles.

In cat peroneus tertius muscles, static gamma axons were prepared in groups of three to four according to the conduction velocity of their axons (fast, intermediate, or slow). Effects of stimulating these groups (at 20, 30, and 50 Hz) on spindle ensemble discharges during sinusoidal stretch (peak-to-peak amplitude, 0.5 mm; frequency linearly increasing from 0.5 to 8 Hz in 10 s) were compared. Ensemble discharges were obtained by digital treatment of the discharges in afferent fibers from all the spindles in peroneus tertius as recorded from the muscle nerve. Stimulation of each group prevented ensemble discharges from falling to very low levels during shortening phases. However, this effect was clearly larger when the group of fast-conducting axons was stimulated. In view of the known effects of the activation of bag(2) and chain fibers (either separately or together) on single primary ending discharges during comparable sinusoidal stretches, this stronger effect supports the view that static gamma axons with faster conduction velocities are more likely to supply more bag(2) fibers than slower ones. Possibly the proportions of bag(2) and chain fibers activated during motor activity are determined by a recruitment of static gamma motoneurons related to their size.

Comparison of static fusimotor innervation in cat peroneus tertius and longus muscles.

Static fusimotor innervation was compared in cat peroneus longus and tertius muscles because the gamma to spindle ratio is considerably higher in the longus (approximately 60 gamma axons for 17 spindles) than in the tertius (approximately 24 gamma axons for 14 spindles). Single gamma axons were identified as static (gamma(s)) by their typical effects on the response of primary ending to ramp stretch. The intrafusal muscle fibers that single gamma(s) axons activated in the spindles they supplied were identified by the features of cross-correlograms between Ia impulses and stimuli, at 100 Hz, and by those of primary ending responses during stimulation at 30 Hz. In each experiment, a large proportion of the gamma population was tested on about nine spindles. A statistical analysis was used to estimate the number of spindles supplied by single gamma(s) axons and the proportion of gamma(s) axons that supply only one spindle among those the stimulation of which had activated either bag2 or chain fibers alone in a single spindle. In peroneus longus, nearly all gamma(s) axons supply one or two spindles, whereas in peroneus tertius, the majority of gamma(s) axons supply from three to six spindles. The proportion of nonspecifically distributed gamma(s) axons, i.e., of axons that supply both bag2 fibers and chain fibers either in the same or in different spindles, is much lower (56%) in the longus than in the tertius (83%) as previously observed on a population of gammas axons that supplied from three to six spindles. Correspondingly, the proportion of specific axons is much higher in the longus (44%) than in the tertius (17%). In none of the two muscles was a strict relationship observed between the conduction velocity of gamma(s) axons and their intrafusal distribution (specific bag2, specific chain fibers, nonspecific). However, gamma(s) supplying bag2 fibers either specifically or in combination with chain fibers tended to have faster conduction velocities, which suggests that, in various motor acts, the proportion of activated bag2 and chain fibers may be related to the proportions of activated fast and slow gamma(s) axons.

Functional consequences of bag2 and chain fiber coactivation by static gamma-axons in cat spindles.

A study of the distribution in cat peroneus tertius spindles of 42 single static gamma-axons was recently carried out with a physiological method for identifying the intrafusal muscle fibers supplied by single gamma-axons. It was found that 35 axons (83%) supplied both slow-contracting bag2 fibers and fast-contracting chain fibers. The distribution of these axons generally varied from one spindle to another among all the spindles that each of them supplied (bag2 and chain fibers together, bag2 alone, chains alone). To find some functional consequences of this coactivation, responses of primary endings to sinusoidal stretch of the muscle (amplitude 0.5-1 mm, frequency linearly increasing from 0.6 to 8-9 Hz in 12 s) were recorded at different average muscle lengths (0.5, 1.0, and 1.5 mm shorter than maximal physiological length) in nembutalized cats during repetitive stimulation at 10, 20, and 30 Hz of single gamma-axons previously shown to supply bag2 and chain fibers in the spindles bearing the primary endings. These responses were compared with responses elicited in passive spindles and during activation of either bag2 fibers or chain fibers alone. Several records of discharge frequency were averaged. During stimulation at 30 Hz of gamma-axons coactivating bag2 and chain fibers, the averaged discharge of primary endings became continuous (that is, without interruption during each shortening phase as occurs in passive spindles) over the whole range of stretch frequencies. The modulation of the discharge was roughly sinusoidal, with an amplitude that increased with the stretch frequency. Stimulation at 30 Hz of gamma-axons activating bag2 fibers alone elicited a modulation of comparable shape and amplitude but only in the range of sinusoidal stretch from 0.6 to 3-4 Hz. Stimulation at 30 Hz of gamma-axons activating chain fibers alone elicited for each cycle in the range of 0.6 to 5-6 Hz a distorted modulation of large amplitude with a minimal frequency close to that of the stimulation. The average muscle length did not significantly influence these various responses. In summary, the coactivation of bag2 and chain fibers, at presumed physiological frequencies, enables primary endings to continuously signal changes of length over a large range of stretch velocities independently of the average muscle length.

Characteristic properties of superficial lumbrical spindles in the cat hind limb, related to their bag1 fibres.

Superficial lumbrical spindles of the cat show several important differences as compared with the classic picture of the spindle, derived principally from tenuissimus. Thus long chain fibres occur in a high proportion of superficial lumbrical spindles, and there is great individual variation in the number of bag fibres present. This study is especially concerned with the morphology and relationships of the bag1, fibre, which are again highly unusual. Most notably, bag1 fibres were almost always the longest intrafusal fibres, typically exceeding bag2 fibres by about 2 mm in total length. They also commonly exhibited the greatest diameter, especially in the extracapsular region where, uniquely, they normally increased in size on leaving the capsule, often becoming similar to extrafusal fibres in this respect. In a few spindles where there was only a single bag fibre this was more often a bag1 than is the case for similar spindles in tenuissimus, and in tandem-linked spindles the continuous bag fibres were more often bag1 than bag2. These results contribute to growing evidence that different muscles exhibit characteristic features of the structure and innervation of their proprioceptors, which may reflect adaptations to local control requirements.

Primary and secondary afferent discharges from the same spindle during chain fibre contraction in cat tenuissimus muscle.

Pairs of Ia and II afferent fibres supplying primary and secondary endings lying in the same tenuissimus spindles were prepared in barbiturate anaesthetized cats in order to compare the variability in the rhythm of discharge of the two endings during responses elicited by the contraction of different intrafusal muscle fibres, especially by chain fibres. In these spindles, the intrafusal muscle fibres supplied by single static gamma-axons were identified with a recently developed technique based on the types of primary ending activation observed during gamma stimulation at 30 and 100 stimuli/s. The responses of the secondary endings to contraction of chain fibres either alone or with bag2 fibres were smaller and much more regular than the responses of the primary endings lying in the same spindles. This difference is probably related to the position of secondary endings along the intrafusal muscle fibres and to the mechanical properties of the muscle fibre regions on which the terminals lie. The dynamic properties of the encoding site of primary afferent impulses probably contribute to the difference. The different degrees of variability observed among secondary ending responses elicited either by chain fibres alone or by chain and bag2 fibres are not related to the type of activated intrafusal muscle fibres.

Distribution of static gamma axons in cat peroneus tertius spindles determined by exclusively physiological criteria.

1. The intrafusal muscle fiber(s) activated in cat peroneus tertius spindles by single static gamma (gamma s) axons were identified by exclusively physiological criteria based on the different contractile properties of chain and bag2 fibers. 2. The identification rested both on the features of primary ending discharges observed during gamma s electrical stimulation at a rate of 30 pulses per second (stimulation at 30/s) and on cross-correlograms constructed during stimulation at 100/s. Three types of primary ending activation could be distinguished. 3. Type F (fast) activations are characterized, at 30/s, by either a 1-to-1 driving or a very irregular increase in firing arising from a level close to the frequency of stimulation and by the presence in cross-correlograms of significant peaks. They are ascribed to chain fibers whose contractions, at 30/s, present large oscillations and, at 100/s, are still incompletely fused. 4. Type S (slow) activations are characterized, at 30/s, by a sustained and generally regular increase in firing and by the absence of significant peaks in cross-correlograms constructed during stimulation at 100/s. They are ascribed to bag2 fibers whose contractions are nearly fused at 30/s and completely fused beyond 60-70/s. 5. Type M (mixed) activations are characterized, at 30/s, by an irregular increase of discharge above a level distinctly higher than the frequency of stimulation and by the presence of significant peaks in cross-correlograms. They are ascribed to the coactivation of chain and bag2 fibers for two reasons: first, they have some features of both type F and type S activations; and second, they are readily reproduced by stimulating together two axons supplying the same spindle, one exerting a type F activation, the other a type S activation. 6. In seven experiments the distribution of 42 single gamma s axons was determined by observing the type of activation they exerted on several spindles (from 3 to 6). Thirty-five axons (83%) were classified "nonspecific" because the type of activation (F, S, or M) varied from one spindle to the other. Seven axons (17%) were classified "specific" because the type of activation was the same in all spindles: either type F for five axons (12%) or type S for two axons (5%). A statistical analysis of the distribution of all activations showed that the proportions of specific axons were not significantly different from those predicted by chance.

Incidence of non-driving excitation of Ia afferents during ramp frequency stimulation of static gamma-axons in cat hindlimbs.

1. The aim of this investigation was to identify static gamma-axons which do not drive any Ia afferents at any stimulus frequency in any spindle which they supply, and to determine their occurrence in various hindlimb muscles (peroneus tertius, brevis, longus and tenuissimus). 2. Ia responses to static gamma stimulation were classified as 'non-driven' when the discharge did not follow the stimulation frequency, or its subharmonics, at any time during a linear increase in stimulus frequency up to 150 Hz lasting 2-3 s, and when tested at two muscle lengths--except in the tenuissimus muscle. In almost all experiments, cross-correlograms were used in addition to evaluate the percentage of these 'non-driven' responses in which a time-locking of discharge to stimulus pulses was obscured by irregularity of the Ia discharge. 3. In 104 spindles, out of 347 responses to stimulation of single static gamma-axons 332 (93%) could be characterized, and of these, 57% (183) were of the non-driven type. The mean number of static gamma effects characterized per spindle was 4.1 (fourteen experiments). In the large majority of spindles (79%, 82 out of 104) at least one response was of the non-driven type. 4. Of the static gamma-axons studied 16% were called 'non driving' ('ndr' gamma s-axons) because they elicited non-driven effects, and since they had the same qualitative effect consistently in all spindles whose discharge was modulated by stimulating them they were called specific 'ndr' axons. If axons with non-driven effects, but acting on one spindle were included in the 'non-driving' category the proportion was 23%. Of spindles tested 63% were innervated by at least one 'ndr' axon. 5. Absence of Ia driving during ramp frequency stimulation of gamma s-axons has been equated with selective bag2 contraction. All the non-driven responses identified in this study cannot be attributed to exclusive bag2 involvement because the total number of 'ndr' responses was too high. In fact, in the isolated spindle preparation bag2 and chain co-contraction were shown to elicit non-driven responses, so chain contraction is not detected reliably in all experimental conditions. Possibly chain fibre contraction is sometimes too weak to dominate the response, or can be of a non-driving character.(ABSTRACT TRUNCATED AT 400 WORDS)

Comparison of skeleto-fusimotor innervation in cat peroneus brevis and peroneus tertius muscles.

1. The skeleto-fusimotor or beta innervation was compared in cat peroneus brevis and peroneus tertius muscles, which differ in their composition of fatigue-resistant motor units; the slow (S) units predominate in brevis and the fast units (FR) in tertius. 2. In four brevis muscles, of thirty-four beta-axons (from a total of 114 axons supplying extrafusal muscle fibres) twenty-nine were dynamic (beta D) and only five static (beta S). In contrast, in three tertius muscles, of twenty-five beta-axons (from a total of 82 axons) twelve were static and thirteen dynamic. 3. In a population of thirty-five brevis and thirty tertius spindles, the proportion of beta D-innervated spindles was greater in the brevis (68.5%) than in the tertius (50%) whereas that of beta S-innervated spindles was greater in the tertius (40%) than in the brevis (17.1%). In a population of thirty-two brevis and twenty-seven tertius spindles in which the presence of bag1 fibres was deduced from the existence of a dynamic innervation, the proportion of spindles innervated by beta D-axons was 80% in the brevis and 62% in the tertius. 4. In both muscles, the number of beta D effects was greater than that of beta S effects. beta S-axons were rarely found to supply more than one spindle whereas beta D-axons supplying more than one spindle (up to four) were common. Spindles were often coinnervated by beta D- and beta S-axons.(ABSTRACT TRUNCATED AT 250 WORDS)

Summation of tension in motor units of the soleus muscle of the cat.

In the cat soleus muscle which is exclusively composed of slow motor units the discrepancy between the sum of individual tensions and the tension on combined stimulation of several motor units was found to be much less than previously reported for slow motor units of peroneus longus. In peroneus the tension on combined stimulation was systematically larger than the value predicted from the sum of individual tensions. For both muscles it was possible to reduce the difference between observed and expected values by comparing the tension on combined stimulation with the sum of tensions, not of single motor units, but of groups of units. It is concluded that whenever tension is measured for single motor units, especially slow units in mixed muscles, the values obtained may be modified by frictional forces. The size of the effect appears to vary from one preparation to the next.

Individual differences in multiple-bag spindles of cat superficial lumbrical muscles.

A total of 791 spindle poles was analysed with regard to intrafusal fibre composition in the first and second superficial lumbrical muscles from the right and left hindfeet of 9 male and 5 female adult cats. Bag and chain muscle fibres were identified by their myofibrillar ATPase staining profile in the B region, after either acid or alkaline preincubation. A high proportion of the spindle pole population (43.2%) was observed to contain three or more (up to 5) bag fibres; those poles were classified as multiple-bag spindle poles. In the 334 muscle spindles in which both poles were studied, 42 bag fibres (12.6%) were found to be of the 'mixed' type, that is a fibre in which the two poles differ in their ATPase staining profile (either bag1/bag2 or bag/chain). The variability of the intrafusal fibre content observed in spindles of these muscles has been studied in relation to individual characteristics such as sex, weight and side of the animal. In general, multiple-bag spindles are more frequent in male than in female cats and in right as compared to left side muscles. Nearly all 'mixed' bag intrafusal fibres (38 out of 42) were observed in spindles containing 3 or more bag fibres. In 3-bag spindles the proportion of 'mixed' bag spindles is approximately the same in male and female cats. The ratio of 'dynamic' (mean polar bag1 content) to 'static' (mean polar bag2 plus chain fibre content) intrafusal effectors per muscle tends to increase in spindles of right side muscles and to decrease in the heaviest animals. The quantitative and qualitative differences in fibre content of spindles observed in first lumbrical muscles of different animals suggest that the spindle fibre composition, especially that of the 'dynamic' bag1 fibre, may be related to individual predetermined and/or acquired factors.

Changes in muscle stiffness produced by motor units of different types in peroneus longus muscle of cat.

1. The effects of maximal tetanic contractions of varying numbers of motor units of the same type [slow (S), fast fatigue-resistant (FR), or fast fatigable (FF)] on the mechanical responses to muscle stretch were studied in the peroneus longus muscle of anesthetized cats. 2. Two types of stiffness measurements were made: 1) an average stiffness, defined as the tension change from the beginning to end of a 0.5-mm ramp stretch; and 2) a dynamic stiffness, defined as the ratio of peak-to-peak tension to amplitude of a maintained 85-microns sinusoidal stretch at frequencies of 10-80 Hz. 3. Contractions of slow and fast units elicited different increases in average stiffness. Type S units, although developing much smaller tetanic tensions than fast ones, produced a resistance to stretch comparable with or greater than that of fast units developing much higher tensions. 4. For comparable tetanic tensions, slow units also elicited a greater dynamic stiffness than fast units. During sinusoidal stretch, changes in muscle tension led changes in muscle length during contraction of S units, but the reverse was observed for frequencies 30-50 Hz during contraction of FF units. This suggests that the latter perform oscillatory work on the driving apparatus. 5. Type S units, whose low-threshold motoneurons are the first to be recruited, appear well adapted to play a role in posture and in slow movements because of the resistance they offer to forces tending to change joint position or to oppose the progression of slow movements.

Proportion of fatigue-resistant motor units in hindlimb muscles of cat and their relation to axonal conduction velocity.

1. A study of motor units to hindlimb muscles of cat has been made, with as complete a sample as possible of the motor axons to an individual muscle. In single experiments as much as 95% of the motor supply to a muscle has been examined. 2. The following muscles have been studied: peroneus brevis, peroneus tertius, peroneus longus, plantaris, gastrocnemius medialis, soleus, tenuissimus and lumbricalis superficialis. 3. Units were identified as slow resistant (S), fast resistant (FR), fast fatigable (FF) and fast intermediate (FI). The proportion of various motor unit types differs from one muscle to another. There is also some variation in the proportions to a given muscle from one animal to another. With the exceptions of soleus, which is entirely slow resistant, and gastrocnemius, which has relatively fewer resistant units, most muscles contain 60% or more of resistant (S and FR) units. 4. The conduction velocity ranges of FF, FR and FI units overlapped. There was little overlap between the conduction velocity ranges of these F units and of S units. 5. In individual experiments there was a strong and significant positive correlation between the logarithm of maximal tetanic tension and axonal conduction velocity in S and in S+FR units. In terms of contractile response the total fatigue-resistant population appeared to be a continuum. The correlation coefficient between maximal tetanic tension and conduction velocity was also high in the totality of units of all types, although within the FF group there appeared to be little or no correlation. In pooled data there was much more scatter and these relations were less clear. This resulted largely from differences in the ranges of axonal conduction velocity for a given motor unit type from one animal to another. 6. There was a highly significant negative correlation between isometric twitch contraction time and axonal conduction velocity in individual experiments. This relationship could also be seen, but less clearly, in pooled data. 7. The possible bases for these relationships are discussed.

High incidence of long-chain fibers in spindles of cat superficial lumbrical muscles.

Forty-two complete spindle poles of cat superficial lumbrical muscles were analyzed with particular regard to the length and the diameter of intrafusal fiber types. Poles were reconstructed from serial transverse sections of fresh-frozen muscles. The staining module, which was repeated throughout the whole muscle, comprised sections treated for glycogen detection and sections treated for detecting myofibrillar ATPase activity after preincubation at three different pH's (see METHODS). The identification of intrafusal fiber types was essentially based on the ATPase activity of the B region of the intrafusal fibers. Long-chain fibers, i.e., chain fibers that have at least one pole that extends by more than one millimeter beyond the end of the spindle capsule (6), were very commonly observed. Of 42 spindle poles analyzed, 30 (71%) contained at least one long-chain fiber (one in 17 spindle poles, 2 in 11 poles, and 3 in 2 poles). Of 246 poles of chain fibers, 45 (18%) were "long". In four spindles, in which both poles could be completely examined, 10 long-chain fibers were observed. In eight of these, only one pole was long; the opposite pole ended either intracapsularly or at a short distance outside the capsule. Since long-chain fiber poles, presently considered to be among the effectors of static skeletofusimotor (beta) axons, are present in a large proportion of muscle spindles of lumbrical muscles, it would be of particular interest to reevaluate the beta-supply of these muscles by physiological methods.

[Summation of maximal tetanic tensions developed by slow or fast motor units of the peroneus longus muscle in the cat]. / Sommation de tensions tétaniques maximales développées par des unités motrices lentes ou rapides du muscle long péronier du chat.

The maximal tetanic tension developed by the simultaneous contraction of several slow motor units (measured tension) is always greater than the sum of the tensions developed by each unit individually (cumulated tension). Generally, the same holds true for fast units but, for measured tensions of same value, the difference between measured tensions and cumulated tensions is much smaller. It is sometimes negligible or even negative.

[Comparison of the increase of stiffness produced during sinusoidal stretch of the long fibula muscle in the cat by the contraction of slow or fast motor units]. / Comparaison de l'augmentation de raideur produite au cours de l'étirement sinusoïdal du muscle long péronier du chat par la contraction d'unités motrices lentes ou rapides.

The increase in stiffness of cat peroneus longus muscle elicited by the contraction of homogeneous groups of motor units (slow, fast fatigue-resistant and fast fatigable) was measured during sinusoidal stretches (20-80 Hz) of small amplitude (40-100 micron). For comparable tetanic tensions, slow units increase muscle stiffness more than fast unit.

[Glycogen depletion produced in intrafusal nuclear bag muscle fibers by brief large-amplitude muscle stretches]. / Déplétion glycogénique produite dans les fibres musculaires intrafusales à sac nucléaire par des étirements musculaires brefs de forte amplitude.

The glycogen content of the three types of intrafusal muscle fibre was studied with histochemical techniques in cat muscle spindles of superficial lumbrical muscles after a very large number of brief large stretches. Zones of glycogen depletion were observed in a high proportion of nuclear bag fibres, notably in bag 1 fibres, but not in chain fibres. These observations suggest that stretching of bag fibres by itself may activate these fibres.

Fusimotor after-effects on responses of primary endings to test dynamic stimuli in cat muscle spindles.

Conditioning stimulation of individual dynamic fusimotor axons, either gamma or beta, leaves after-effects which enhance the responses of primary endings to test stimulation of the same axon applied during slow ramp stretch. These after-effects have a long duration, persisting well over 5 min, but are abolished by stretch of large amplitude. The dynamic after-effects also enhance frequencygrams elicited by low-frequency repetitive stimulation during slow ramp stretch, causing single stimuli to become much more effective. When several dynamic axons to the same spindle are isolated, conditioning stimulation of one leaves an after-effect to test stimulation of itself and of all other dynamic axons. When two dynamic axons are used for conditioning stimulation, facilitation or occlusion can be demonstrated in their interaction, indicating that they converge on the same intrafusal element. Dynamic after-effects persist during background static fusimotor activity of considerable amplitude, suggesting that static and dynamic actions are quite independent. Dynamic after-effects appear to result from residual changes in the bag fibre, probably from a persistent increase in the number of cross-bridges between thick and thin filaments. These after-effects produce a large increase in the response of primary endings to dynamic fusimotor activity and probably have an important functional role.

Effects of stretch on dynamic fusimotor after-effects in cat muscle spindles.

Conditioning stimulation of dynamic fusimotor axons leaves persistent after-effects which increase the responses of primary endings to test dynamic stimuli. Such after-effects are abolished by muscle stretch. Destruction of these after-effects depends on the following. (a) Amplitude of stretch: with symmetrical triangular stretches of moderate velocity, an extension of soleus by 4-5 mm totally abolishes the after-effects. Lesser stretches cause a graded reduction. (b) Velocity of relaxation: for a given amplitude of stretch there is greater destruction of after-effects when it is followed by a slow rate of relaxation than after rapid relaxation. (c) After-effects tested late in ramp stretch are more resistant to destruction by stretch than those increasing test dynamic responses early in ramp stretch. Stretch itself produces after-effects which enhance test responses to dynamic but not to static fusimotor stimulation. Interactions between conditioning dynamic stimulation and stretch suggest that both these effects occur in the same intrafusal elements, the bag fibres.

Glycogen depletion elicited in tenuissimus intrafusal muscle fibres by stimulation of static gamma-axons in the cat.

In this study the experimental conditions used to elicit glycogen depletion in tenuissimus intrafusal muscle fibres were different from those used by Barker, Emonet-Dénand, Harker, Jami & Laporte (1976): the tenuissimus was left in situ; several (4-6) static gamma-axons were stimulated together; the blood flow through the muscle was not reduced during the periods of gamma stimulation except in two experiments; very much longer periods (up to 9 h) of intermittent stimulation by bursts at 50-500/s were used. Bag1 and bag2 fibres were identified by their different ATPase activities in the B region. In two experiments with normal circulation, test responses of several primary endings to short periods of stimulation at 50-100/s were still very strong after stimulation of several static gamma-axons for 5 and 9 h, respectively. Glycogen depletion was observed in a large number of chain and bag2 poles but in only one of nineteen bag1 poles examined. In two other experiments with normal circulation, there was a very pronounced reduction of the test responses after stimulation of several static gamma-axons for 7 and 9 h, respectively. Out of twenty-four bag1 poles examined, nineteen exhibited zones of depletion. In an experiment in which stimulation was conducted as in Barker et al. (1976), i.e. with reduction of muscle blood flow during 1 min periods of stimulation at 50-100/s, the primary endings still gave a strong response after fifteen periods of stimulation in contrast with the marked 'fatigue' that was constantly observed in the former study. No depleted intrafusal fibres were found in the spindles of this muscle. In a last experiment, after an initial pattern of stimulation similar to that described above, the new pattern of stimulation, but with periodical reduction of blood flow, was applied, leading to a 'fatigue' of the test responses in 2 h. In the spindles of this muscle three out of ten bag1 poles were depleted. The variability of glycogen depletion in bag1 fibres appears to be linked to the degree of spindle 'fatigue' which may develop after static gamma stimulation. It seems that in 'fatigued' spindles some factor or factors liberated by the contraction of neighbouring fibres may deplete glycogen in bag1 fibres by a non-neural mechanism. When, in spite of a prolonged stimulation of static gamma-axons, no fatigue of the test responses develops, zones of depletion in bag1 fibres--possibly of neural origin--are very rare, although a large proportion of bag2 and chain fibres are depleted.

Observations on the effects on spindle primary endings of the stimulation at low frequency of dynamic beta-axons.

Stimulation of single dynamic beta-axons at presumed physiological frequencies (10-30/s) significantly increases the sensitivity of spindle primary endings of cat peroneus brevis muscle to ramp and sinusoidal stretches. This action becomes greater as the amplitude of stretch and the initial muscle length is increased. Stimulation at low frequency of two dynamic beta-axons supplying the same spindle greatly enhances the effect elicited by each axon individually.