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.

     

Characteristic curves of the dynamic response of primary muscle spindle endings with and without gamma stimulation

1. The characteristic curves of the dynamic response of primary muscle spindle endings (i.e. the dependence of dynamic response on the stretch volocity) have been recorded with and without gamma stimulation in 75 muscle spindles of the cats' M. tibialis anterior. They can be described by a power function DR=a · v _r. ⁿ
2. Under dynamic fusimotor stimulation the prefactor ‘a’ increases and the exponent ‘n’ decreases; under static gamma stimulation vice versa.
3. The characteristic curves with and without gamma stimulations of a single spindle are intersecting, because of the change of the exponent ‘n’.
4. The dyamic response under dynamic gamma stimulation usually increases, but we observed a decrease, too. Under static gamma stimulation the dynamic response usually decreases.
5. In the following we looked on the gamma stimulations statistically. We had 47 dynamic gamma stimulations. Here we observed a separation into an upper and a lower group of the characteristic curves, which proved to be statistically significant, according to their effects on the course of the characteristic curves and the velocity sensitivity.
6. The lower group of the dynamic gamma stimulations and all 24 static gamma stimulations do not differ significantly i) from the group of characteristic curves without gamma stimulation and ii) in the absolute course of the characteristic curves and in the velocity sensitivity under gamma stimulation.
7. The upper group of dynamic gamma stimulations is the only one to cause significantly strong effects on the course of the characteristic curves and velocity sensitivity.
8. The upper group of dynamic gamma fibre stimulations extends the measurable velocity range to very small stretch velocities, which otherwise do not cause a measurable dynamic response.

     

Organization of group I activated cells in the main and external cuneate nuclei of the cat: Identification of muscle receptors

Summary Extracellular recording was made from 77 primary afferent fibres, 106 cells in the external cuneate nucleus, and 60 cells in the main cuneate nucleus, all activated by slowly adapting muscle stretch receptors. The nature of the muscle receptors responsible for the activation was determined by various types of receptor stimulation.Primary group I afferents from muscle spindles and tendon organs in distal forelimb muscles showed complete overlap of conduction velocities and thresholds to electrical stimulation. Both types of group I afferents as well as group II muscle spindle afferents were shown to ascend through the dorsal funiculus to the level of the cuneate nuclei.Three groups of cells were identified in the external cuneate nucleus, activated by group I muscle spindle afferents, tendon organ afferents and group II muscle spindle afferents, respectively.Almost all group I activated cells in the main cuneate nucleus, including all 34 cells identified as cuneo-thalamic relay cells, received their afferent input from muscle spindle afferents. Three cells were activated by tendon organ afferents.     

Acetylcholine-sensitivity and local regenerative activity in denervated frog slow muscle fibres

1. The distribution of acetylcholine (ACh)-sensitive membrane areas was determined in 11–75 days denervated slow muscle fibres ofRana temporaria by iontophoretic application of acetylcholine. The fibres were also stimulated directly, and their electrical activity was recorded with an intra- and an extracellular electrode.
2. During the first two weeks following denervation the size of the ACh-sensitive fibre surface was similar to that of normal slow fibres, but a spreading out of ACh-sensitivity occurred between the 13th and 20th day.
3. The slow fibre membrane did not become homogeneously ACh-sensitive; even after long periods of denervation large local sensitivity gradients could be observed. Throughout the denervation period maximum values of ACh-sensitivity were in the same range as in normal slow fibres.
4. Action potentials were fully developed when ACh-sensitivity started to spread out. Extracellularly recorded inward currents varied in amplitude along the fibre surface, and either one or two peaks were observed in individual fibres.
5. The spatial relationship between inward current peaks and peaks of ACh-sensitivity was investigated in 12 fibres. Fifteen inward current peaks were located at distances of 30–640 μm from points of maximum ACh-sensitivity; only once did the centers of ACh-sensitivity and excitability coincide.
6. It is concluded that Na channels are incorporated into nonjunctional membrane areas of denervated slow fibres; this process preceds the incorporation of ACh-receptors by approximately one week.

     

A comparative analysis of the encapsulated end-organs of mammalian skeletal muscles and of their sensory nerve endings

The encapsulated sensory endings of mammalian skeletal muscles are all mechanoreceptors. At the most basic functional level they serve as length sensors (muscle spindle primary and secondary endings), tension sensors (tendon organs), and pressure or vibration sensors (lamellated corpuscles). At a higher functional level, the differing roles of individual muscles in, for example, postural adjustment and locomotion might be expected to be reflected in characteristic complements of the various end‐organs, their sensory endings and afferent nerve fibres. This has previously been demonstrated with regard to the number of muscle‐spindle capsules; however, information on the other types of end‐organ, as well as the complements of primary and secondary endings of the spindles themselves, is sporadic and inconclusive regarding their comparative provision in different muscles. Our general conclusion that muscle‐specific variability in the provision of encapsulated sensory endings does exist demonstrates the necessity for the acquisition of more data of this type if we are to understand the underlying adaptive relationships between motor control and the structure and function of skeletal muscle. The present quantitative and comparative analysis of encapsulated muscle afferents is based on teased, silver‐impregnated preparations. We begin with a statistical analysis of the number and distribution of muscle‐spindle afferents in hind‐limb muscles of the cat, particularly tenuissimus. We show that: (i) taking account of the necessity for at least one primary ending to be present, muscles differ significantly in the mean number of additional afferents per spindle capsule; (ii) the frequency of occurrence of spindles with different sensory complements is consistent with a stochastic, rather than deterministic, developmental process; and (iii) notwithstanding the previous finding, there is a differential distribution of spindles intramuscularly such that the more complex ones tend to be located closer to the main divisions of the nerve. Next, based on a sample of tendon organs from several hind‐foot muscles of the cat, we demonstrate the existence in at least a large proportion of tendon organs of a structural substrate to account for multiple spike‐initiation sites and pacemaker switching, namely the distribution of sensory terminals supplied by the different first‐order branches of the Ib afferent to separate, parallel, tendinous compartments of individual tendon organs. We then show that the numbers of spindles, tendon organs and paciniform corpuscles vary independently in a sample of (mainly) hind‐foot muscles of the cat. Grouping muscles by anatomical region in the cat indicated the existence of a gradual proximo‐distal decline in the overall average size of the afferent complement of muscle spindles from axial through hind limb to intrinsic foot muscles, but with considerable muscle‐specific variability. Finally, we present some comparative data on muscle‐spindle afferent complements of rat, rabbit and guinea pig, one particularly notable feature being the high incidence of multiple primary endings in the rat.     

Changes in PAD patterns of group I muscle afferents after a peripheral nerve crush

In the anesthetized cat we have analyzed the changes in primary afferent depolarization (PAD) evoked in single muscle spindle and tendon organ afferents at different times after their axons were crushed in the periphery and allowed to regenerate. Medial gastrocnemius (MG) afferents were depolarized by stimulation of group I fibers in the posterior biceps and semitendinosus nerve (PBSt), as soon as 2 weeks after crushing their axons in the periphery, in some cases before they could be activated by physiological stimulation of muscle receptors. Two to twelve weeks after crushing the MG nerve, stimulation of the PBSt produced PAD in all MG fibers reconnected with presumed muscle spindles and tendon organs. The mean amplitude of the PAD elicited in afferent fibers reconnected with muscle spindles was increased relative to values obtained from Ia fibers in intact (control) preparations, but remained essentially the same in fibers reconnected with tendon organs. Quite unexpectedly, we found that, between 2 and 12 weeks after crushing the MG nerve, stimulation of the bulbar reticular formation (RF) produced PAD in most afferent fibers reconnected with muscle spindle afferents. The mean amplitude of the PAD elicited in these fibers was significantly increased relative to the PAD elicited in muscle spindle afferents from intact preparations (from 0.08–0.4 to 0.47-0.34 mV). A substantial recovery was observed between 6 months and 2.5 years after the peripheral nerve injury. Stimulation of the sural (SU) nerve produced practically no PAD in muscle spindles from intact preparations, and this remained so in those afferents reconnected with muscle spindles impaled 2–12 weeks after the nerve crush. The mean amplitude of the PAD produced in afferent fibers reconnected with tendon organs by stimulation of the PBSt nerve and of the bulbar RF remained essentially the same as the PAD elicited in intact afferents. However, SU nerve stimulation produced a larger PAD in afferents reconnected with tendon organs 2–12 weeks after the nerve crush (mean PAD changed from 0.05-0.04 to 0.32-0.17 mV). The results obtained indicate that the PAD patterns of the afferent fibers reconnected with muscle spindle and tendon organ afferents are changed after crushing their axons in the periphery: stimulation of the bulbar RF appears to produce larger PAD in fibers reconnected with muscle spindles, and stimulation of cutaneous afferents produces larger PAD in fibers reconnected with tendon organs. It is suggested that these alterations in the patterns of PAD of muscle afferents result from central changes in the balance of excitatory and inhibitory influences acting on the segmental pathways mediating the PAD. Although the functional role of these changes has not been established, they may reflect compensatory changes aimed to adjust information arising from damaged afferents.     

The afferent innervation of two infrahyoid muscles of the cat

1. The afferent innervation of the straplike muscles of the infrahyoid region were investigated in two ways. The morphology of spindles and counts of tendon organs were investigated by the gold chloride technique in ten muscles. Spindle counts were made in forty pairs of thyrohyoid and infrahyoid muscles. De-efferenting of the nerves to these muscles was done in three cats and the calibre spectra of the afferent innervation investigated. These were compared with the total counts of fibres in intact nerves.2. In the thyrohyoid, spindles are frequently absent. No tendon organs were seen. In the large infrahyoid (combined sternohyoid and sternothyroid), spindle counts varied from 0 to 20 and the mean spindle count per gram of muscle was 3.5. A maximum of five tendon organs were seen in the muscle. Both spindle and tendon organ counts are low when compared with a limb muscle of similar weight and size.3. In the infrahyoid muscle complex spindles were about equal in number to simple spindles.4. Counts of spindles in the infrahyoid muscle in families of three or more siblings suggest that some families of kittens tend to have higher spindle counts than other families.5. The afferent innervation of the two muscles varied between 21 and 42% of the total fibre population and the fibre diameter spectrum is in keeping with the low counts of encapsulated endings.     

Batrachotoxin protects sodium channels from the blocking action of oenanthotoxin

1. The blocking action of oenanthotoxin (OETX) and butanol on Na⁺ channels was studied in voltage clamp experiments on single myelinated nerve fibres treated by batrachotoxin (BTX).
2. OETX (40 μM) blocked Na⁺ currents through normal channels but did not affect significantly the BTX modified Na⁺ current.
3. BTX removed the depolarization-induced charge immobilization and slowed down significantly the OFF charge movement. However, the maximum charge displaced, as well as the kinetics of the ON charge movement during a strong membrane depolarization, remained unchanged.
4. OETX blocked the charge movement in normal Na⁺ channels but did not affect noticeably the charge movement modified by BTX.
5. BTX did not modify the K⁺ currents and did not protect them from the blocking action of OETX.
6. Butanol (0.01–0.1 M) decreased almost identically and reversibly both normal and BTX-modified Na⁺ currents.
7. It is concluded that binding of BTX to its receptor protects the Na⁺ channel from interaction with OETX but left it accessible to butanol.

     

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.     

Physiological properties of muscle spindles in dorsal neck muscles of the cat.

1. Single-fiber recording was used to examine the properties of 107 spindle endings in cat biventer cervicis (BC) and complexus (CM) muscles. Responses of receptors were examined following muscle contraction and ramp and hold stretch. Twenty-two endings in splenius (SP) were also examined, but their responses could not be quantitated because the anatomy of SP prevented the application of appropriate stretches. 2. Conduction velocitites of spindle afferents ranged from 13 to 90 m/s. Endings with primary response patterns usually had faster conduction velocities than secondary endings, but there was overlap in the conduction velocity ranges of the two subgroups. 3. Most neck spindle afferents could be classified as either primary or secondary by a constellation of physiological criteria including dynamic response pattern, dynamic index, and variability of resting discharge frequency. However, 22 of 107 endings from BC and CM had responses with characteristics intermediate between primary and secondary responses. The possible sources of these characteristics are discussed. 4. Despite the similarity in properties between spindles of different neck muscles, the length sensitivities of CM spindles were high compared to those of BC spindles. CM spindles showed length-related modulation of firing frequency over a more restricted range of initial muscle lengths than did BC spindles. 5. Eight Golgi tendon organs (GTO) were identified by their characteristics responses. Conduction velocities obtained for five GTO afferent nerves ranged from 50 to 67 m/s. Recordings were also made from receptros in deep muscles surrounding the vertebrae. These receptors had properties characteristic of muscle spindles.     

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.     

Resting discharge of human muscle spindles is not modulated by increases in sympathetic drive

There is evidence in experimental animals that, in addition to receiving fusimotor drive, muscle spindles are subject to modulation by the sympathetic nervous system. We examined the validity of this idea in human subjects by recording from muscle spindles in the relaxed ankle and toe extensor muscles during a strong and sustained physiological activation of muscle sympathetic outflow. Unitary recordings were made from 20 primary and 17 secondary muscle spindle afferents via a tungsten microelectrode inserted percutaneously into the peroneal nerve in 10 awake, healthy subjects seated with the legs supported in the extended position. ECG, blood pressure, respiration and calf circumference were also recorded. The majority of the muscle spindles were spontaneously active at rest; a background discharge was induced in four silent spindles by vibrating the tendon. A sustained increase in muscle vasoconstrictor activity, an increase in calf volume and a fall in pulse pressure were produced by subjects performing a 30-40 s maximal inspiratory breath-hold. Despite this strong increase in muscle sympathetic outflow no significant changes occurred in the discharge of either primary or secondary muscle spindle afferents, measured as a change in mean frequency and variability over sequential 5 s epochs and compared with the preceding period of rest. Strong chemoreceptor-driven sympathetic bursts during sustained expiratory breath-holds also failed to modulate the firing of 14 spindle endings. We conclude that a sustained, physiological increase in muscle sympathetic activity causes no detectable change in muscle spindle firing, lending no support to the concept that the sympathetic nervous system can influence the sensitivity of human muscle spindles directly.     

Properties of cat neck muscle spindles and their excitation by succinylcholine

Summary The sensitivity to sinusoidal stretching and small-amplitude vibration, and the variability of the resting discharge rate of de-efferented muscle spindles in the neck extensor muscle biventer cervicis (b.c.) of the cat have been studied. The effects of intra-arterial infusion of succinylcholine (SCh) on the response of the receptors to ramp stretches of the muscle were also determined. When activated by SCh, one group of afferents showed only a slow facilitation of their discharge rate, similar to that of spindle secondary sensory endings in hind-limb muscles. A second group of afferents developed a large dynamic response and a marked increase in their static discharge rate and were presumed to originate in normal b₁b₂c spindles in the b.c. muscle. A third group of afferents developed only a marked increase in static discharge, without potentiation of the dynamic response, suggesting an origin in the b₂c units of tandem spindles which are relatively common in the neck muscles. On the basis of their passive characteristics alone, afferents from b₁b₂c units could not be readily distinguished from those from b₂c units. The characteristics of these receptors, and their differences from the well-studied hind-limb spindle afferents, are discussed.     

The influence of extrafusal muscle activity on discharge patterns of primary muscle spindle endings

The influence of extrafusal muscle activity in anaemically decerebrate cats upon discharge patterns of primary spindle endings was ascertained by simultaneously recording spike trains from several Ia afferents and muscle tension fluctuations of the triceps-surae muscle. 1. Tension fluctuations were averaged with respect to spikes from primary endings yielding tension "trajectories" of specific shape for each spindle and probably reflecting frequently recurring mechanical events in the spindles' surroundings. 2. Spindles situated in close vicinity and influenced by similar mechanical events as evidenced by similar average tension trajectories are correlated in their discharge patterns to a degree depending on the strength of their mechanical coupling. 3. The modulation of spindle discharge frequency in response to average tension changes at the muscle tendon is very different in amplitude for different spindles; this response may show a high sensitivity. It is usually phase advanced by 90-180 degrees with respect to the "internal length" changes; between spindles ther may be phase differences of up to 180 degrees. 4. It is concluded that primary endings react very sensitively to local extrafusal events. The CNS receives much more accurate information about these events in the correlation of several Ia afferents than in the discharge of a single fibre.     

Termination patterns of identified group II and III afferent fibres from deep tissues in the spinal cord of the cat

In chloralose-anaesthetized cats, the impulse activity of single afferent fibres supplying receptors in the deep tissues of the hindlimb (fasciae, muscles, ligaments, joint capsules) was recorded using micropipettes filled with a solution of horseradish peroxidase. Only myelinated fibres with conduction velocities up to 40 m/s (Group III and Group II units) were studied, i.e. fast conducting afferent fibres from muscle spindles and tendon organs were excluded. The fibres were functionally characterized with the use of mechanical stimuli such as local pressure and joint movements. The results show that a relationship exists between the functional properties of a given afferent unit and the location of its terminals in the spinal cord. Since the conduction velocity and hence the diameter of the fibres was similar in all the units studied, these factors appear not to be of importance for determining the pattern of spinal termination. Out of 84 units, 42 were classified as high-threshold mechanosensitive, 26 as low-threshold mechanosensitive, and 16 as secondary endings from muscle spindles. Following physiological identification the fibres were ionophoretically injected with horseradish peroxidase and their trajectory in the white and gray matter of the spinal cord visualized histologically with diaminobenzidine. High-threshold mechanosensitive units took a lateral course in the posterior funiculus and usually did not bifurcate. They exhibited two different patterns of spinal termination, one being characterized by terminal arborizations in both lamina I and deeper laminae (mostly IV/V), the other one by an exclusive projection to lamina I. Low-threshold mechanosensitive units often showed a bifurcation in the posterior funiculus and did not have a uniform termination pattern. The main areas of termination were lamina II and laminae IV-VI. The slowly conducting secondary endings from muscle spindles projected mainly to laminae VI and VII with additional collaterals entering the ventral horn. They thus had a termination pattern similar to that reported for fast conducting afferent fibres (above 50 m/s) from muscle spindle secondary endings. With the exception of one high-threshold mechanosensitive unit none of the stained fibres possessed terminal arborization and boutons in lamina III. It is concluded that different types of Group II and III primary afferent fibres from deep tissues exhibit different patterns of spinal termination.(ABSTRACT TRUNCATED AT 400 WORDS)     

Reabsorption of monocarboxylic acids in the proximal tubule of the rat kidney

1. Renal handling of pig-and rat-insulin was studied in the isolated perfused rat kidney.
2. Metabolic clearance rates of both pig- and rat-insulin excceded GFR.
3. Peritubular uptake of pig-insulin accounted for 13%, of rat-insulin for 31% of the total metabolic clearance.
4. The nonfiltering kidney does not remove insulin from the peritubular circulation.
5. Metabolic clearance rates of pig- and rat-insulin are directly related to GFR.
6. The filtration process seems to be necessary for the uptake of insulin at the peritubular site.

     

Evidence of the co-activation of α-motoneurones and static γ-motoneurones of the sartorius medialis muscle during locomotion in the thalamic cat

Summary The activity in and efferent axon populations and in group I and group II afferent fibre populations innervating a flexor muscle, the sartorius medialis, was observed during spontaneous locomotor movements in the thalamic cat. Multi-unit discharges of each kind of fibre were obtained by electronic sorting of the action potentials from the overall activity of a thin, intact branch of the sartorius medialis nerve. The following results were obtained: (1) The -motoneurones have a phasic behaviour characterized by a single discharge period during the hip flexion (swing phase of the step-cycle). (2) The -motoneurones are co-activated with the homonymous -motoneurones. (3) Between rhythmic and discharges, i.e. during the hip extension (stance phase of the step cycle), both - and -motoneurones were normally silent. However, in 5 out of 17 experiments, a few units of the population fired at very low frequency. (4) Two observations indicate that the -motoneurones that are co-activated with the -motoneurones by central locomotor commands are predominantly of the static type. In actual locomotion, the rhythmic fusimotor discharges over-compensate the depressor effect on the firing rate of the group II afferents of the unloading of muscle spindles by the active shortening of the parent muscle. In fictive locomotion, when the transmission of the excitation is blocked by selective curarization in alpha skeleto-motor junctions alone, the rhythmic fusimotor discharges elicit in-phase modulations not only of the group I but also of the group II fibres. The group II afferent population consists almost entirely of fibres arising from spindle secondary endings which are located primarily on intrafusal muscle fibres whose contraction is exclusively controlled by static fusimotor motoneurones. In the two experimental circumstances, the analysis of the group I fibre discharge does not allow to decide whether dynamic motoneurones are firing or silent during rhythmic discharge. (5) The group I and group II afferent discharges during the step-cycle showed two frequency peaks, one static-fusimotor dependent while the contracting muscle shortened during the hip flexion (swing) phase, the other length-change dependent while the relaxed muscle was rapidly stretched during the first part of the hip extension (stance) phase. Then, during the second part of hip extension when the muscle was slowly stretched in the absence of fusimotor drive, the firing rate of the spindle afférents decreased to a low level. The spindle sensory endings during the extension phase showed low dynamic and static responsiveness like deefferented spindles. (6) The results obtained in sartorius medialis (flexor) muscle are discussed in comparison with the results previously obtained in gastrocnemii (extensor) muscles (Bessou et al. 1986). The consequences of the predominant activation of the static or dynamic fusimotor system in functionally different muscles are considered with respect to the proprioceptive or motor role of musclespindles during muscle contraction.     

Segmental and supraspinal control of synaptic effectiveness of functionally identified muscle afferents in the cat

The present investigation documents the patterns of primary afferent depolarization (PAD) of single, functionally identified muscle afferents from the medial gastrocnemius nerve in the intact, anesthetized cat. Classification of the impaled muscle afferents as from muscle spindles or from tendon organs was made according to several criteria, which comprised measurement of conduction velocity and electrical threshold of the peripheral axons, and the maximal frequency followed by the afferent fibers during vibration, as well as the changes in discharge frequency during longitudinal stretch, the projection of the afferent fiber to the motor pool, and, in unparalyzed preparations, the changes in afferent activity during a muscle twitch. In confirmation of a previous study, we found that most muscle spindle afferents (46.1–66.6%, depending on the combination of criteria utilized for receptor classification) had a type A PAD pattern. That is, they were depolarized by stimulation of group I fibers of the posterior biceps and semitendinosus (PBSt) nerve, but not by stimulation of cutaneous nerves (sural and superficial peroneus) or the bulbar reticular formation (RF), which in many cases inhibited the PBSt-induced PAD. In addition, we found a significant fraction of muscle spindle primaries that were depolarized by stimulation of group I PBSt fibers and also by stimulation of the bulbar RF. Stimulation of cutaneous nerves produced PAD in 9.1–31.2% of these fibers (type B PAD pattern) and no PAD in 8.2–15.4% (type C PAD pattern). In contrast to muscle spindle afferents, only the 7.7–15.4% of fibers from tendon organs had a type A PAD pattern, 23–46.1% had a type B and 50–61.5% a type C PAD pattern. These observations suggest that the neuronal circuitry involved in the control of the synaptic effectiveness of muscle spindles and tendon organs is subjected to excitatory as well as to inhibitory influences from cutaneous and reticulospinal fibers. As shown in the accompanying paper, the balance between excitation and inhibition is not fixed, but can be changed by crushing the afferent axons in the peripheral nerve and allowing subsequent reconnection of these afferent fibers with muscle receptors.     

Static γ-motoneurones couple group Ia and II afferents of single muscle spindles in anaesthetised and decerebrate cats

Ideas about the functions of static γ-motoneurones are based on the responses of primary and secondary endings to electrical stimulation of single static γ-axons, usually at high frequencies. We compared these effects with the actions of spontaneously active γ-motoneurones. In anaesthetised cats, afferents and efferents were recorded in intramuscular nerve branches to single muscle spindles. The occurrence of γ-spikes, identified by a spike shape recognition system, was linked to video-taped contractions of type-identified intrafusal fibres in the dissected muscle spindles. When some static γ-motoneurones were active at low frequency (< 15 Hz) they coupled the firing of group Ia and II afferents. Activity of other static γ-motoneurones which tensed the intrafusal fibres appeared to enhance this effect. Under these conditions the secondary ending responded at shorter latency than the primary ending. In another series of experiments on decerebrate cats, responses of primary and secondary endings of single muscle spindles to activation of γ-motoneurones by natural stimuli were compared with their responses to electrical stimulation of single γ-axons supplying the same spindle. Electrical stimulation mimicked the natural actions of γ-motoneurones on either the primary or the secondary ending, but not on both together. However, γ-activity evoked by natural stimuli coupled the firing of afferents with the muscle at constant length, and also when it was stretched. Analysis showed that the timing and tightness of this coupling determined the degree of summation of excitatory postsynaptic potentials (EPSPs) evoked by each afferent in α-motoneurones and interneurones contacted by terminals of both endings, and thus the degree of facilitation of reflex actions of group II afferents.     

Actions of afferent impulses from muscle receptors on cerebellar Purkyně cells. II. Responses to muscle contraction: Effects mediated via the climbing fiber pathway

Summary The effect of isometric contraction of the gastrocnemius-soleus muscle group was investigated on the responses of 79 cerebellar Purkyn cells (P-cells) in the decerebrate cat. Forty three percent of the P-cells tested were responsive to contraction via a pathway that acted by way of climbing fibres. The latency of these responses was long (mean 60 msec following ventral root stimulation) and variable. It was concluded that the responses were attributable to Group Ib afferent input. An excitatory action of the contraction on spindle afferent fibres was excluded because the responses were: a) evoked by submaximal contractions produced by weak single shock stimulation of the ventral root; b) negligible following isotonic contractions; and, c) augmented when a stretch was superimposed on the rising phase of the twitch.The effects of both muscle contraction and muscle vibration were studied for many of the P-cells in order to investigate the convergence from afferents of both spindle and tendon organ origin. Few P-cells had evidence of a convergent input from Group Ia, II, and Ib fibres. In contrast convergence from Group II and Group Ib input was quite common. With some cells, there appeared to be an input from only Group Ib or Group II fibres.