Testing the classification of static gamma axons using different patterns of random stimulation.

The possibility of using randomly generated stimulus intervals (with a Poisson distribution) to identify the type(s) of intrafusal fiber activated by the stimulation of single static gamma axons was tested in Peroneus tertius muscle spindles of anesthetized cats. Three patterns of random stimulation with different values of mean intervals [20 +/- 4. 47, 30 +/- 8.94, and 40 +/- 8.94 (SD) ms] were used. Single static gamma axons activating, in single spindles, either the bag2 fiber alone or the chain fibers alone or both types of intrafusal fiber were prepared. Responses of spindle primary endings elicited by the stimulation of gamma axons were recorded from Ia fibers in cut dorsal root filaments. Cross-correlograms between stimuli and spikes of the primary ending responses, autocorrelograms, interval histograms of responses, and stimulations were built. The characteristics of cross-correlograms were found to be related not only to the type of intrafusal muscle fibers activated but also to the parameters of the stimulation. Moreover some cross-correlograms with similar characteristics were produced by the activation of different intrafusal muscle fibers. It also was observed that, whatever the type of intrafusal muscle fiber activated, cross-correlograms could exhibit oscillations after an initial peak, provided the extent in frequency of the primary ending response was small; these oscillations arise in part from the autocorrelation of the primary ending responses. Therefore, cross-correlograms obtained during random stimulation of static gamma axons cannot be used for unequivocally identifying the type(s) of intrafusal muscle fiber these axons supply.     

On the subdivision of static and dynamic fusimotor actions on the primary ending of the cat muscle spindle.

1. Using large ramp and triangular stretches a survey has been made of the effect of stimulating single gamma fusimotor fibres on primary endings of muscle spindles in the peroneus brevis to see whether 'intermediate' types of fusimotor action could be recognized, falling between the well known static and dynamic types. 2. Responses were classified into six groups, as detailed on pp. 844-846, ranging from apparently 'pure' dynamic action (category I) to apparently 'pure' static action (category IV). Models for a putative mixed action were produced by combining the stimulation of a static and of a dynamic fibre to the same spindle. The clearest sign of static action was firing on the releasing phase of the stretch. The essential sign of dynamic action, which survived combination with the more dominant static action, was the low adaptive decay of firing with a time constant of about 0-5 sec that occurs on the plateau of the ramp stretch. 3. Out of 153 responses, each elicited from a primary ending on stimulation of a single fusimotor fibre, 67% were apparently 'pure' examples of dynamic and static action. The remaining 33% of responses were to some degree suggestive of an admixture, in various proportions, of static and dynamic actions. For only 18% of them was there firm indication of such admixture. 4. When a given fibre was tested on more than one ending then, with one exception out of thirty-six instances, its action always proved to be either predominantly static or predominantly dynamic. There was no special tendency for an axon with a mixed action on one spindle to have a similarly mixed action on other endings so that individual fusimotor fibres were best classified as static or dynamic without intermediate grades. 5. Simultaneous stimulation of two fusimotor fibres eliciting apparently 'pure static and dynamic actions, could mimic all the intermediate types of action. 6. The results are discussed in relation to recent studies, especially those based on glycogen depletion. It was concluded that dynamic action arises from activation of the bag1 intrafusal muscle fibre, and that static action arises from the bag2 and chain fibres, whether acting individually or collaboratively. The intermediate actions are suggested to arise from an overlap of motor innervation to contrasting types of intrafusal muscle fibre. 7. On the basis of effects on the regularity of the afferent discharge the findings support the view that a given static action axon can innervate bag2 and chain fibres in various proportions in different spindles, so that they do not provide separable effector pathways. 8. Responses to large amplitude sinusoidal stretching were also studied in relation to our classification.     

The Influence of Bag2 and Chain Intrafusal Muscle Fibres on Secondary Spindle Afferents in the Cat

Static γ-motor activity is strongly modulated by a particular phase relationship to the cyclic movements of locomotion, and this has a profound effect on the firing patterns of muscle spindle afferents. Whilst primary afferents are affected by both static and dynamic γ-motor output, secondary afferents are affected significantly only by the static system acting via the intrafusal bag₂ and chain fibres. It is therefore important to know how fluctuating patterns of static γ-motor activity affect secondary afferents and to relate this to the actions of bag₂ and chain fibres. We have studied the action of single static γ axons on secondary afferents in cat hindlimb muscles. Various physiological methods were explored to identify which of the intrafusal muscle fibres were being activated in each case, including the use of random stimulation and ramp frequency stimulation. The effects were also recorded of 1 Hz sinusoidally frequency-modulated γ-axon stimuli and the amplitude and phase of the resulting afferent modulation related to the involvement of the bag₂ and chain fibres. It was found that bag₂ fibres are effective in biasing the secondary discharge, but their modulating action is relatively weak and involves a marked phase lag. Chain fibres acting alone cause strong modulation with very little phase lag. Mixed bag₂ and chain-fibre action is most effective in modulating afferent discharge and causes intermediate values of phase lag. The results are discussed in relation to the control of natural movements and it is concluded that an important function of the static γ motor system is to provide a signal to sum algebraically with the length-related signal. The results do not suggest that it could also usefully control stretch sensitivity.     

Motor and sensory innervation of muscle spindles in the neonatal rat

Summary Neural and muscular elements of three muscle spindles from the soleus muscles of 4-day-old rats were reconstructed by electron microscopy of skip-serial transverse ultrathin sections. Each spindle contained four encapsulated intrafusal fibers, including a minimum of one bag₁, one bag₂ and one chain fiber. The fibers were innervated by unmyelinated motor and sensory axons. The primary and secondary afferents approached the spindles as single axons and terminated on the central region of the intrafusal fibers. Single profiles of terminal axons occupied the sites of sensory neuromuscular junctions, similar to adult sensory endings. No morphological features suggested retraction of afferents from 4-day postnatal spindles. Motor axons approached spindles tightly packed in bundles of 5–20 axons and terminated in the juxtaequatorial and polar regions of both bag and chain fibers. Multiple profiles of terminal axons were visible for each intrafusal motor ending. More motor axons innervated 4-day postnatal spindles and a greater number of axon terminals were visible in immature intrafusal motor endings than in adult spindles. The data suggest that postnatal maturation of motor innervation to intrafusal fibers involves the elimination of supernumerary motor nerve inputs. Synapse elimination in the development of the fusimotor system might represent a mechanism whereby individual axons adjust the number of spindles they innervate.     

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.     

Spindle secondary ending responses elicited by stimulation of static fusimotor axons

1. Indirect evidence on the time course of intrafusal events associated with stimulation of a static fusimotor axon was obtained by analysis of the responses from muscle spindle secondary endings of the cat.2. In many instances, a single impulse in a single static fusimotor axon to a muscle spindle evoked a transient increase in discharge of the secondary ending the duration of which ranged from 30 to 60 msec with a rising phase of 6-20 msec. Increasing the length of the muscle increased the slope of the rising phase and augmented the magnitude of the response.3. Different static fusimotor axons produced different effects upon secondary endings. A single impulse in some fusimotor axons did not produce an effect while repetitive stimulation produced a smooth increase in discharge frequency. When efficacious, a single fusimotor impulse evoked approximately a similar duration of increased discharge from a secondary ending and a primary ending of the same muscle spindle. The relative magnitude of effects of a given fusimotor axon on primary and secondary endings of a spindle varied.4. During partial curarization the changes in the discharges of a secondary ending evoked by stimulation of a single static fusimotor axon occurred in a step-like fashion.5. These results are interpreted as supporting the idea that static fusimotor axons induce twitch-like contractions of intrafusal elements (nuclear chain) through the intermediary of propagated action potentials.     

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.     

Glycogen-depletion method of intrafusal distribution of gamma-axons that increase sensitivity of spindle secondary endings

1. The glycogen-depletion method was used in cat tenuissimus muscles to investigate whether the action of static gamma-axons that increase the sensitivity of spindle secondary endings to muscle length is associated with a specific pattern of distribution of these axons to intrafusal muscle fibers. 2. In each experiment, a single gamma-axon acting on a secondary ending of a precisely located spindle was repetitively stimulated, and subsequently the intrafusal muscle fibers of that spindle were examined for glycogen depletion. 3. The gamma-axons whose repetitive stimulation increased the length sensitivity of secondary endings depleted all the chain fibers in one or both poles of the spindle, with the bag fibers being inconstantly involved. The gamma-axons whose stimulation did not exert this effect produced much more restricted patterns of glycogen depletion. 4. The length sensitivities of two secondary endings belonging to the same spindle were similarly affected by a single gamma-axon, whereas a gamma-axon could increase the sensitivity of a secondary ending without altering that of the primary ending of the same spindle. 5. The action exerted by single gamma-axons on secondary endings appears related to their intrafusal distribution since enhancement of the secondary endings sensitivity was observed only when all the chain fibers of at least one spindle pole were activated. 6. If several static gamma-axons supplying a given spindle are firing together, an increase in the length sensitivity of the secondary endings of that spindle can be expected since probably all chain fibers are activated.     

The regularity of primary and secondary muscle spindle afferent discharges

1. The patterns of nerve impulses in the afferent fibres from muscle spindles have been studied using the soleus muscle of the decerebrate cat. Impulses from up to five single units were recorded simultaneously on magnetic tape, while the muscle was stretched to a series of different lengths. Various statistics were later determined by computer analysis.2. After the ventral roots were cut to eliminate any motor outflow to the muscle spindles, both primary and secondary spindle endings discharged very regularly. At frequencies around 30 impulses/sec the coefficient of variation of the interspike interval distributions had a mean value of only 0.02 for the secondary endings and 0.058 for the primary endings. The values obtained for the two kinds of ending did not overlap.3. When the ventral roots were intact, the ;spontaneous' fusimotor activity considerably increased the variability of both kinds of endings. Secondary endings still discharged much more regularly than primary endings, even when the fusimotor activity increased the frequency of firing equally for the two kinds of endings. At frequencies around 30/sec the average coefficient of variation of the interval distributions was then 0.064 for the secondary endings and 0.25 for the primary endings.4. When the ventral roots were intact there was usually an inverse relation between the values of successive interspike intervals. The first serial correlation coefficient often had values down to - 0.6 for both kinds of ending. Higher order serial correlation coefficients were also computed.5. Approximate calculations, based on the variability observed when the ventral roots were intact, suggested that when the length of the muscle was constant an observer analysing a 1 sec period of discharge from a single primary ending would only be able to distinguish about six different lengths of the muscle. The corresponding figure for a secondary ending was twenty-five lengths.6. The increase in variability with fusimotor activity, and the pattern of serial correlations, were probably caused by static fusimotor fibres firing at rates below the fusion frequency of the intrafusal muscle fibres that they supply.     

Action of static and dynamic fusimotor fibres on secondary endings of cat's spindles

1. The effects exerted on secondary endings by repetitive stimulation of single static or single dynamic fusimotor fibres were studied in nine tenuissimus spindles in cats.2. The discharge of each secondary ending was recorded simultaneously with the discharge of the primary ending belonging to the same spindle.3. All the nineteen static fusimotor fibres studied activated secondary endings.4. Of eight dynamic fusimotor fibres, seven had no action on secondary endings. One dynamic fibre activated an atypical secondary ending which displayed some phasic sensitivity.5. No difference in conduction velocity was found between static and dynamic fibres.6. The implications of these observations for the mechanism of action of static and dynamic fusimotor fibres are discussed.     

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.     

The effects of suxamethonium and acetylcholine on the behaviour of cat muscle spindles during dynamics stretching, and during fusimotor stimulation

1. The responses of primary and secondary afferent fibres from muscle spindles in cat soleus were studied during constant velocity stretching.2. Intravenous Suxamethonium (SCh) caused a large increase in the response of the primary afferents to the dynamic phase of stretching, and a smaller increase in their response to static extension. The effects of SCh were similar to the effects of dynamic fusimotor stimulation.3. Increasing doses of SCh increased the response of primary afferents to dynamic stretching up to a point, but a peak discharge frequency was encountered beyond which the afferent fibre could not be induced to discharge.4. Suxamethonium increased the response of the secondary afferent fibres by a smaller amount than the primaries, and in particular caused a smaller increase in the response to dynamic extension.5. Acetylcholine given by close arterial injection had an effect similar to the effect of SCh.6. The effects of static and dynamic fusimotor stimulation on the response of primary afferents summed with the effects of small doses of SCh. When large amounts of SCh were used fusimotor stimulation sometimes had no further effect on the afferent discharge. It was not possible to say whether the fusimotor activity was then inhibited, or submerged in the SCh activity.7. The actions of SCh and of acetylcholine emphasized the differences in response of primary and secondary afferent endings to dynamic stretching. The use of these drugs enabled us to classify fibres of intermediate conduction velocity.8. Suxamethonium is known to activate slow muscle fibre systems with distributed nerve endings. The similarity between dynamic fusimotor activity and the effect of SCh suggests that the dyanimic fusimotor fibres act on slow intrafusal muscle fibres through multiple distributed endings.     

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.     

Innervation of developing intrafusal muscle fibers in the rat

The chronology of development of spindle neural elements was examined by electron microscopy in fetal and neonatal rats. The three types of intrafusal muscle fiber of spindles from the soleus muscle acquired sensory and motor innervation in the same sequence as they formed—bag₂, bag₁, and chain. Both the primary and secondary afferents contacted developing spindles before day 20 of gestation. Sensory endings were present on myoblasts, myotubes, and myofibers in all intrafusal bundles regardless of age. The basic features of the sensory innervation—first-order branching of the parent axon, separation of the primary and secondary sensory regions, and location of both primary and secondary endings beneath the basal lamina of the intrafusal fibers—were all established by the fourth postnatal day. Cross-terminals, sensory terminals shared by more than one intrafusal fiber, were more numerous at all developmental stages than in mature spindles. No afferents to immature spindles were supernumerary, and no sensory axons appeared to retract from terminations on intrafusal fibers. The earliest motor axons contacted spindles on the 20th day of gestation or shortly afterward. More motor axons supplied the immature spindles, and a greater number of axon terminals were visible at immature intrafusal motor endings than in adult spindles; hence, retraction of supernumerary motor axons accompanies maturation of the fusimotor system analogous to that observed during the maturation of the skeletomotor system. Motor endings were observed only on the relatively mature myofibers; intrafusal myoblasts and myotubes lacked motor innervation in all age groups. This independence of the early stages of intrafusal fiber assembly from motor innervation may reflect a special inherent myogenic potential of intrafusal myotubes or may stem from the innervation of spindles by sensory axons.     

Effects of changes in preoptic temperature on stretch response of muscle spindle endings in the cat's soleus muscle

1. The effects of changing the temperature in the preoptic region on the stretch responses of primary and secondary endings of the muscle spindle in the soleus muscle was investigated in urethane anesthetized cats. The local temperature of the preoptic region was controlled by implanting water perfused thermodes (32–42.5°C).
2. A standard ramp and hold stretch stimulus was repeatedly applied to the soleus muscle during changes of preoptic temperature from normal to hypo-or hyperthermic values. During each stretch the instantaneous firing rate was recorded and the static firing frequency and the dynamic index were electronically determined according to Crowe and Matthews (1964).
3. Of a total of 76 investigated muscle spindle afferents, the stretch responses of 49 (65%) were altered by changing preoptic temperature. The static stretch response was increased in all cases during cooling, whereas during preoptic heating the static stretch response increased in about half of the afferents and decreased in the rest. Increase in the static stretch response during both cooling and heating was usually combined with the appearence of spontaneous discharges at rest.
4. With regard to the static stretch response, primary and secondary muscle spindle endings responded similarly to cooling and heating.
5. Two types of preoptic temperature effects on the stretch responses of primary endings were observed. In the majority of afferents only the static stretch response was augmented during cooling, whereas the peak response was little changed resulting in a decrease of the dynamic index (‘static’ response type). In a minority of primary afferents the effect of preoptic cooling consisted of a greater increase of the peak response than of the static stretch response. i.e., the dynamic index also increased (‘dynamic-static’ response type). In secondary muscle spindle endings only the static type of response to preoptic cooling was observed.
6. It is concluded that changes of preoptic temperature influence the stretch response of muscle spindle afferents mainly by altering the activity of the static fusimotor innervation, its activation by preoptic cooling being the most consistent finding. In a minority of muscle spindles the dynamic fusimotor innervation appears to be additionally activated by preoptic cooling.

     

Effects of temperature on the discharges of muscle spindles and tendon organs

1. In anaesthetized cats the effects of temperature on the nervous outflow from skeletal muscle via thick myelinated afferent fibres were studied. Single unit recordings were made from afferents of muscle spindles and tendon organs during slow and fast temperature changes of the medial gastrocnemius muscle which was deefferented by ventral root section and prestretched to a tension of 100 p.
2. Group I afferent units from muscle spindles were activated by warming and depressed by cooling, the effect of warming being much more pronounced than that of cooling. Afferents from secondary spindle endings with a high background discharge behaved similar to Ia fibres, whereas those with a low initial discharge rate showed an activation by cooling and a depression (mostly to cessation of firing) by warming. The discharges of group I afferents from tendon organs varied; an activation by warming was the most frequently observed reaction.
3. Some of the afferents from muscle spindles and tendon organs showed signs of a dynamic sensitivity to thermal stimulation, but in general the dynamic component in the responses to temperature changes was only small.
4. The results suggest that the afferent outflow via thick myelinated fibres from a resting, moderately prestretched muscle strongly depends on temperature. At raised intramuscular temperatures (about 42°C) the nervous outflow is characterized by an increased activity in all of the I a and many of the I b afferents, while the majority of group II spindle afferents will be depressed. In contrast, in a cold muscle (about 29°C) the nervous outflow via afferents from primary spindle endings will be reduced, while the net activity from secondary spindle endings will be increased and no marked changes are expected to occur in the discharges of I b fibres.

     

Correlation of the dynamic behaviour of deefferented primary muscle spindle endings with their static behaviour

20 deefferented primary muscle spindle endings, 8 afferents from the extensor digitorum longus muscle (EDL) and 12 from the tibialis anterior muscle (TA), were investigated in 12 anesthetized cats. Dynamic responses of muscle spindles to ramp stretches were correlated with their static responses. The variation in the state and dynamic behaviour was found to be interdependent. The correlation coefficients were higher for EDL spindles than for TA spindles. The correlation coefficients depended upon the definition of the dynamic responses.     

The relative sensitivity to vibration of muscle receptors of the cat

1. Longitudinal vibration was applied to the de-efferented soleus muscle of anaesthetized cats while recording the discharge of single afferent fibres from the proprioceptors within the muscle. Conditions were defined under which vibration can be used to excite selectively the primary endings of muscle spindles without exciting the secondary endings of muscle spindles or Golgi tendon organs.2. Frequencies of vibration of 100-500 c/s were used. The maximum amplitude of vibration which the vibrator could produce fell with increasing frequency; it was 250 mu (peak to peak) for 100 c/s and 20 mu for 500 c/s.3. Primary endings of muscle spindles were very sensitive to vibration. Most could be ;driven' to discharge one impulse for each cycle of vibration over the whole of the above range of frequencies, provided the initial tension was moderate (20-200 g wt.). The amplitude of vibration required to produce driving usually varied by less than a factor of two over the whole range of frequencies. The most sensitive endings could be driven by vibrations of below 10 mu amplitude.4. Stimulation of single fusimotor fibres, whether static or dynamic fusimotor fibres, increased the sensitivity of primary endings to vibration. Contraction of the main muscle, produced by stimulating alpha motor fibres, reduced the sensitivity of primary endings even when fusimotor fibres were also being stimulated.5. The secondary endings were very insensitive to longitudinal vibration and with the amplitudes available not one of twenty-five endings could be driven at 150 c/s or above; one ending could be driven at 100 c/s by vibration of 250 mu amplitude. Stimulation of single fusimotor fibres, probably all of which were static fusimotor fibres, made them slightly more sensitive to vibration but none of them approached the sensitivity of the primary endings.6. The Golgi tendon organs were as insensitive as the secondary endings when the muscle was not contracting and none could be driven at any frequency in spite of quite high tensions in the muscle. However, when the muscle was made to contract by stimulating alpha fibres in ventral root filaments the tendon organs became appreciably more sensitive, the degree of sensitization increasing approximately with the strength of the contraction. They never became as sensitive as the primary endings, and with the amplitudes of vibration available none was driven at frequencies of over 250 c/s.7. When the amplitude of vibration was somewhat below that required to produce driving of an ending it still produced some increase in its mean frequency of discharge. However, amplitudes of vibration of 25-50 mu applied to a non-contracting muscle, whether with or without fusimotor stimulation, produced driving of nearly all primary endings without any significant increase in the mean frequency of firing of secondary endings or Golgi tendon organs. Such vibration can therefore be used as a specific stimulus for the primary endings in order to investigate the central effects or repetitive discharge of the Ia afferent fibres from them.8. Experiments on endings in the peroneus longus muscle showed that these behaved similarly to those in soleus.     

Differential presynaptic inhibition of actions of group II afferents in di- and polysynaptic pathways to feline motoneurones

The aim of this study was to investigate differences in the effects of presynaptic inhibition of transmission from group II muscle afferents to neurones in the dorsal horn and in the intermediate zone and the consequences of these differences for reflex actions of group II afferents upon α-motoneurones. The degree of presynaptic inhibition was estimated from the degree of depression of monosynaptic components of population EPSPs (field potentials) evoked by group II muscle afferents in deeply anaesthetized cats. The decrease in the area of field potentials was considerably larger and longer lasting in the intermediate zone, where they were often obliterated, than in the dorsal horn, where they were reduced to about two-thirds. Presynaptic inhibition of field potentials evoked by other afferents at the same locations was much weaker. Intracellular records from α-motoneurones revealed that short latency EPSPs and IPSPs evoked from group II afferents are considerably reduced by conditioning stimuli that effectively depress intermediate zone field potentials of group II origin. The results of this study lead to the conclusion that strong presynaptic inhibition of transmission to intermediate zone interneurones allows a selective depression of disynaptic actions of group II muscle afferents on α- and γ-motoneurones, mediated by these interneurones, and favours polysynaptic actions of these afferents.     

Effects upon spindle discharges of electrical stimulation of static fusimotor fibers with concomitant application of muscle vibration.

Discharges of single afferent fibers from the primary endings of the soleus muscle spindles were recorded from thin dorsal root filaments in cats anesthetized with urethane and chloralose. The distal cut end of the ventral root was split into fine filaments to obtain functionally single fusimotor fibers. The fusimotor fibers obtained in this study were of the static type. The soleus muscle was sinusoidally stretched at 70 Hz with various amplitude concurrently with 100 Hz electric stimulation of fusimotor fiber. The spindle afferent discharges were analysed by compiling inter-spike interval histograms and cross-correlograms between the afferent spikes and the stimulus pulses applied to the fusimotor fiber. The same analysis was also made between the afferent spikes and peak extensions of muscle yielded by vibratory stimulation. One-third of the fusimotor fibers were capable of driving the spindle afferents. The driving of fusimotor stimulation was replaced by driving by muscle dibration of more than 10 mum amplitude applied concurrently with fusimotor stimulation. The remaining two-thirds of the fusimotor fibers could not drive the spindle afferents. In this case, the driving by muscle vibration was obtained when vibration of more than 5 mum amplitude was applied concurrently with fusimotor stimulation. It was suggested that fusimotor fibers which produced driving of the spindle afferents would terminate on nuclear chain fibers and those not producing driving on nuclear bag fibers, or the latter would terminate relatively distant from the primary ending as compared with the former.