How shunting inhibition affects the discharge of lumbar motoneurones: a dynamic clamp study in anaesthetized cats.

In the present work, dynamic clamp was used to inject a current that mimicked tonic synaptic activity in the soma of cat lumbar motoneurones with a microelectrode. The reversal potential of this current could be set at the resting potential so as to prevent membrane depolarization or hyperpolarization. The only effect of the dynamic clamp was then to elicit a constant and calibrated increase of the motoneurone input conductance. The effect of the resulting shunt was investigated on repetitive discharges elicited by current pulses. Shunting inhibition reduced very substantially the firing frequency in the primary range without changing the slope of the current-frequency curves. The shift of the I-f curve was proportional to the conductance increase imposed by the dynamic clamp and depended on an intrinsic property of the motoneurone that we called the shunt potential. The shunt potential ranged between 11 and 37 mV above the resting potential, indicating that the sensitivity of motoneurones to shunting inhibition was quite variable. The shunt potential was always near or above the action potential voltage threshold. A theoretical model allowed us to interpret these experimental results. The shunt potential was shown to be a weighted time average of membrane voltage. The weighting factor is the phase response function of the neurone that peaks at the end of the interspike interval. The shunt potential indicates whether mixed synaptic inputs have an excitatory or inhibitory effect on the ongoing discharge of the motoneurone.

Positive proprioceptive feedback elicited by isometric contractions of ankle flexors on pretibial motoneurons in cats.

Pretibial flexor motoneurons were recorded intracellularly in anesthetized cats during unfused isometric contractions of a subpopulation of motor units from either tibialis anterior (TA) or extensor digitorum longus (EDL) muscles. The contractions elicited excitatory postsynaptic potentials in 23 of 28 pretibial flexor motoneurons. No effect was observed in the remaining motoneurons. In control experiments, the effects of electrical stimulation of afferents within the TA nerve were investigated to help identify afferents responsible for the contraction-induced positive feedback. This feedback was ascribed to actions of Ia fibers because the pattern of the contraction-induced excitatory potentials was consistent with the known pattern of Ia discharge; in control experiments, electrical stimulation of group I fibers elicited only monosynaptic excitatory potentials; and the distribution of both the contraction-induced positive feedback among motor nuclei as well as the electrically evoked Ia excitatory monosynaptic potentials were restricted to homonymous and synergic motoneurons. Observation of the Ia contraction-induced positive feedback was facilitated by the absence of Ib autogenic inhibition. This contraction-induced Ia excitatory feedback in ankle flexors might either reinforce Ia-induced reflexes when these muscles are lengthened or help to lift the leg over an obstacle.

Quantitative evidence for multiple widespread representations of individual muscles in the cat motor cortex.

We sought to determine why a given muscle appears represented in widespread loci in the motor cortex (MCx). To this end, we microstimulated every 500 microm along medio-lateral rows and recorded the evoked electromyographic (EMG) responses of up to a dozen forelimb muscles of the cat. A consistent finding in all animals studied was that along a given row, distal muscle responses could be elicited from medially situated cortical loci and conversely, proximal muscles responses from laterally situated cortical loci. In many such cases, the evoked EMG responses were such that the largest responses from a distal muscle were obtained by stimulation at a medially situated point and those of a proximal muscle from a laterally situated point. A Spearman correlation analysis showed that there was no correlation between cortical position and where the peak response of a given muscle occurred. These quantitative results strongly support the view that in the forelimb area of the cat MCx there exists widespread 'colonies' of corticospinal neurons with common spinal cord targets.

Cooperation of muscle and cutaneous afferents in the feedback of contraction to peroneal motoneurons.

Peroneal motoneurons were recorded intracellularly in anesthetized cats during sustained submaximal contractions of peroneus brevis muscle (PB) elicited by repetitive electrical stimulation of motor axons in the distal portion of cut ventral root filaments. Mechanical stimulation of the territory innervated by the superficial peroneal nerve (SP) was applied during contraction to assess the influence of afferents from this territory on the contraction-induced excitation of motoneurons. In 21 peroneal motoneurons in which PB contraction evoked excitatory potentials, a stimulation engaging mechanoreceptors located in the skin around toes was found to either enhance (in 12 motoneurons) or reduce (in 9 motoneurons) the contraction-induced excitatory potentials. Among positive effects, six showed simple summation of the responses to each individual stimulus, suggesting a convergence of afferent pathways on motoneurons. In six other motoneurons, complex interactions were observed, as may result from convergence at a premotoneuronal level. Among negative effects, a single instance was observed of inhibitory facilitation, as may result from convergence of cutaneous and muscular, possibly Ib, afferents on inhibitory interneurons. Several pathways, mediating either facilitory or inhibitory influences, are available for cooperation of muscle and cutaneous input, allowing flexibility of motoneuron activation in different tasks.

Effects on peroneal motoneurons of cutaneous afferents activated by mechanical or electrical stimulations.

The postsynaptic potentials elicited in peroneal motoneurons by either mechanical stimulation of cutaneous areas innervated by the superficial peroneal nerve (SP) or repetitive electrical stimulation of SP were compared in anesthetized cats. After denervation of the foot sparing only the territory of SP terminal branches, reproducible mechanical stimulations were applied by pressure on the plantar surface of the toes via a plastic disk attached to a servo-length device, causing a mild compression of toes. This stimulus evoked small but consistent postsynaptic potentials in every peroneal motoneuron. Weak stimuli elicited only excitatory postsynaptic potentials (EPSPs), whereas increase in stimulation strength allowed distinction of three patterns of response. In about one half of the sample, mechanical stimulation or trains of 20/s electric pulses at strengths up to six times the threshold of the most excitable fibers in the nerve evoked only EPSPs. Responses to electrical stimulation appeared with 3-7 ms central latencies, suggesting oligosynaptic pathways. In another, smaller fraction of the sample, inhibitory postsynaptic potentials (IPSPs) appeared with an increase of stimulation strength, and the last fraction showed a mixed pattern of excitation and inhibition. In 24 of 32 motoneurons where electrical and mechanical effects could be compared, the responses were similar, and in 6 others, they changed from pure excitation on mechanical stimulation to mixed on electrical stimulation. With both kinds of stimulation, stronger stimulations were required to evoke inhibitory postsynaptic potentials (IPSPs), which appeared at longer central latencies than EPSPs, indicating longer interneuronal pathways. The similarity of responses to mechanical and electrical stimulation in a majority of peroneal motoneurons suggests that the effects of commonly used electrical stimulation are good predictors of the responses of peroneal motoneurons to natural skin stimulation. The different types of responses to cutaneous afferents from SP territory reflect a complex connectivity allowing modulations of cutaneous reflex responses in various postures and gaits.

Flexible processing of sensory information induced by axo-axonic synapses on afferent fibers.

Recent experiments indicate that afferent information is processed in the intraspinal arborisation of mammalian group I fibres. During muscle contraction, Ib inputs arising from tendon organs are filtered out by presynaptic inhibition after their entry in the spinal cord. This paper reviews the mechanisms by which GABAergic axo-axonic synapses, i.e., the morphological substrate of presynaptic inhibition, exert this filtering effect. Using confocal microscopy, axo-axonic synapses were demonstrated on segmental Ib collaterals. Most synapses were located on short preterminal and terminal branches. Using a simple compartmental model of myelinated axon, the primary afferent depolarisation (PAD), generated by such synapses, was predicted to reduce the amplitude of incoming action potentials by inactivating the sodium current, and this prediction was experimentally verified. A further theoretical work, relying on cable theory, suggests that the electrotonic structure of collaterals and the distribution of axo-axonic synapses allow large PADs (about 10 mV) to develop on some distal branches, which is likely to result in a substantial presynaptic inhibition. In addition, the electrotonic structure of group I collaterals is likely to prevent PAD from spreading to the whole arborisation. Such a non-uniform diffusion of the PAD accounts for differential presynaptic inhibition in intraspinal branches of the same fibre. Altogether, our experimental and theoretical works suggest that axo-axonic synapses can control the selective funnelling of sensory information toward relevant targets specified according to the motor task.

Indications for GABA-immunoreactive axo-axonic contacts on the intraspinal arborization of a Ib fiber in cat: a confocal microscope study.

Confocal microscopy was used to detect GABA-immunoreactive axo-axonic appositions, indicating possible synaptic contacts, on Ib fiber terminals in the lumbosacral spinal cord. A Ib fiber from posterior biceps-semitendinosus muscles was labeled by intra-axonal ejection of tetramethylrhodamine dextran (red), and serial sections of S1-L7 spinal cord segments were processed for GABA immunocytochemistry revealed by fluorescein isothiocynate (green). Appositions between GABA-immunoreactive structures and the labeled fiber appeared as yellow spots because of the presence of both fluorochromes in small volumes (0.3 * 0.3 * 0.5 micrometer(3)) of tissue. These spots were identified as probable axo-axonic contacts when: (1) they were observed in two to four serial confocal planes, indicating that they did not occur by chance; and (2) their sizes, shapes, and locations were similar to those of axo-axonic contacts found on Ia terminals, known to bear presynaptic boutons, and resembled the axo-axonic synapses described in electron microscope studies of Ib boutons in Clarke's column. A total of 59 presumed axo-axonic contacts was observed on two Ib collaterals, representing an estimated 20% of the total complement. In a three-dimensional reconstruction of one collateral, they were mostly located in terminal positions, and some branches bore more contacts than others. Such differential distribution could not result from chance appositions between GABAergic structures and Ib arborization and further supported the identification of axo-axonic contacts. Segmental Ib collaterals bear axo-axonic synapses that might ensure differential funneling of information toward different targets.

Reduction of presynaptic action potentials by PAD: model and experimental study.

A compartmental model of myelinated nerve fiber was used to show that primary afferent depolarization (PAD), as elicited by axo-axonic synapses, reduces the amplitude of propagating action potentials primarily by interfering with ionic current responsible for the spike regeneration. This reduction adds to the effect of the synaptic shunt, increases with the PAD amplitude, and occurs at significant distances from the synaptic zone. PAD transiently enhances the sodium current activation, which partly accounts for the PAD-induced fiber hyperexcitability, and enhances sodium inactivation on a slower time course, thus reducing the amplitude of action potentials. In vivo, intraaxonal recordings from the intraspinal portion of group I afferent fibers were carried out to verify that depolarizations reduced the amplitude of propagating action potentials as predicted by the model. This article suggests PAD might play a major role in presynaptic inhibition.

Heterogeneity of contraction-induced effects in neurons of the cat dorsal spinocerebellar tract.

1. Clarke's column neurons of the dorsal spinocerebellar tract (DSCT) were recorded intracellularly in anaesthetized cats during weak sustained contractions of triceps surae (TS) produced by direct electrical stimulation of the muscle. 2. Of 145 DSCT neurons, 77 (53%) were contraction sensitive suggesting that information about weak contraction of a limited number of muscles is widely distributed among DSCT neurons. Four types of effects were observed in individual neurons during TS contractions. 3. In the first group of 11 DSCT neurons (14% of the contraction-sensitive cells), the effect was excitation persisting throughout the duration of contractions. These responses were ascribed to actions of afferents from contraction-activated tendon organs. 4. In a second group of 15 neurons (20% of the contraction-sensitive cells), quickly declining excitatory potentials were recorded during sustained TS contractions. By analogy with previous observations of contraction-induced effects in motoneurons, the decline of excitation might be explained by contraction-induced presynaptic inhibition of group I afferents in Clarke's column. 5. Declining inhibitions, resembling those previously observed in homonymous and synergic motoneurons, were recorded in 49% of contraction-sensitive DSCT neurons. This appears in keeping with the fact that interneurons mediating Ib inhibition to motoneurons project axon collaterals to DSCT neurons. Presynaptic inhibition of Ib fibres might therefore cause parallel reductions of inhibitory potentials in motoneurons and in DSCT neurons. 6. In a final group of 13 neurons, mixed excitatory and inhibitory effects were observed during TS contractions. Such DSCT neurons might monitor the excitability of Ib interneurons by integration of information about input to and output from these neurons. 7. The non-uniform patterns of DSCT responses to TS contractions suggest complex processing of information on ankle extensor activity in cerebellum. Phasic signalling of contraction onset is observed in many DSCT neurons while others carry messages about duration and strength of contraction.

Contraction-induced excitation in cat peroneal motoneurons.

1. Motoneurons innervating peroneal muscles were recorded intracellularly in anesthetized cats during sustained submaximal isometric contractions of peroneus brevis produced by repetitive electrical stimulation of motor axons in the distal portion of cut ventral root filaments. 2. In contrast with the inhibition previously observed during contractions of gastrocnemius medialis muscle in triceps surae motoneurons, the afferent input generated by peroneus brevis contraction elicited excitatory potentials in nearly all motoneurons supplying peroneus brevis, peroneus tertius, or peroneus longus muscles. 3. We ascribed the contraction-induced excitation of peroneal motoneurons to spindle afferents for two reasons. First, the amplitude of contraction-induced excitatory potentials increased when the ventral root stimulation strength was increased to recruit gamma-axons. Second, with stimulation strengths under gamma-threshold, peroneus brevis contraction still excited peroneal motoneurons, and we obtained evidence that activation of spindles by skeletofusimotor beta-axons could account at least partly for this excitation. 4. The lack of contraction-induced inhibition in peroneal motoneurons and the prevalence of contraction-induced excitation raised the possibility that, in contrast to the usual effects of tendon organ afferents, Ib afferents from peroneus brevis might exert an excitatory influence on homonymous motoneurons. The fact that electrical stimulation of group I afferents in the nerve to peroneus brevis only exceptionally evoked inhibition in peroneal motoneurons would appear compatible with this hypothesis. Furthermore, stimulation of cutaneous afferents, known to facilitate transmission in Ib pathways, only exceptionally revealed a weak contraction-induced inhibition.(ABSTRACT TRUNCATED AT 250 WORDS)

Declining inhibition in ipsi- and contralateral lumbar motoneurons during contractions of an ankle extensor muscle in the cat.

1. Motoneurons of pretibial ankle flexor and knee flexor and extensor muscles were recorded intracellularly in chloralose- or pentobarbitone-anesthetized cats during sustained submaximal contractions of either ipsi- or contralateral gastrocnemius medialis muscle (GM). 2. In a majority of ipsilateral motoneurons, a sustained GM contraction elicited inhibitory potentials that quickly subsided before the end of the contraction. An abrupt increase in contractile force could elicit a new series of inhibitory potentials, which declined again in spite of a maintained force level. 3. Contraction-induced effects were only exceptionally detected in contralateral triceps surae and plantaris motoneurons. In a small number of pretibial flexor and knee flexor and extensor motoneurons, declining inhibitions were observed during sustained contractions of the contralateral GM muscle. 4. At the onset of GM contractions, a variety of motoneurons uniformly receive inhibitory inputs that are quickly filtered out. Although the functional significance of this widespread initial inhibition remains to be elucidated, its rapid decline seems useful to allow subsequent recruitment of motor units as may be required for coordination of posture and movement. 5. Tendon organs are activated during muscle contraction, but it is not certain whether Ib inputs from GM can account for all the effects observed. Contribution of other afferents was considered and tested using a different experimental approach. The companion paper reports observations suggesting that effects elicited by group II afferents may cooperate in the contraction-induced inhibition of motoneurons.

Declining inhibition elicited in cat lumbar motoneurons by repetitive stimulation of group II muscle afferents.

1. The aim of the present experiments was to verify whether group II inputs from gastrocnemius medialis (GM) muscle could elicit declining inhibitions similar to those observed during GM contractions in a variety of lumbar motoneurons of the cat spinal cord. Motoneurons were recorded intracellularly in chloralose- or pentobarbitone-anesthetized preparations during electrical stimulation of GM nerve with repetitive trains. 2. With strengths in the group I range, repetitive stimulation evoked the usual Ia excitation in homonymous motoneurons and excitatory postsynaptic potential (EPSP) amplitudes remained constant throughout the stimulation sequence. In synergic plantaris motoneurons lacking an excitatory connection with Ia afferents from GM, the same stimulation, kept at a constant strength throughout the stimulation sequence, elicited rapidly decreasing inhibitory potentials reminiscent of those evoked by GM contractions. 3. In motoneurons of pretibial flexors, quadriceps, and posterior biceps-semitendinosus, the stimulation strength required to observe declining inhibitions resembling those produced by GM contractions was 4-8 times group I threshold, engaging group II in addition to group I fibers. 4. These results show that input from GM group II plus group I afferents can elicit inhibitory effects in a variety of motoneurons. Such observations support the hypothesis that messages from spindle secondary endings and/or nonspecific muscle receptors activated during contraction might contribute to the widespread inhibition caused by GM contractions. 5. Inasmuch as constant input in group II and group I afferents evoked declining inhibitory potentials, the origin of the decline must be central, which suggests that the rapid reduction of contraction-induced inhibitions also depended on a central mechanism.

Neuromimetic model of a neuronal filter.

Experimental work in cats has shown that a series of afferent impulses from muscle receptors activated during contractions of an ankle extensor elicit declining inhibitory potentials in homonymous and synergic motoneurones. Inhibitory potentials were ascribed to the action of Ib afferents from Golgi tendon organs that are specific contraction-sensitive mechano-receptors. The decline of inhibition was, at least partly, due to presynaptic inhibition acting as a filter of tendon organ information in the spinal cord. In the present work, a computer model of the simplest spinal pathways from Ib fibres to motoneurones was designed. In order to make the model as realistic as possible, the most pertinent of the known functional properties of the neuronal elements were incorporated. Functions simulating primary afferent depolarizations of Ib terminals, i.e. the electrophysiological correlate of presynaptic inhibition, were introduced in the network. Simulations showed that declining inhibitory potentials were computed in the output stage of the network that represented the "motoneurone-like" element. These results support the assumption that the filtering out of Ib inputs is to a great extent due to presynaptic inhibition. The model behaved as expected, suggesting that predictions of the behaviour of neural components in the biological network should be possible upon introduction in the model of other, more complex, spinal pathways from Ib fibres to motoneurones.

Depolarization of Ib afferent axons in the cat spinal cord during homonymous muscle contraction.

1. Intra-axonal records from the intraspinal course of Ib and Ia afferent fibres innervating the gastrocnemius medialis muscle were obtained in chloralose or Nembutal-anaesthetized cats during submaximal contractions of the muscle. 2. Afferent fibres in continuity with their muscle of origin were functionally identified by their responses to muscle stretch or contraction. 3. In six out of eight Ib afferents, primary depolarizations (PADs) were recorded during contraction. They were independent of the presence of orthodromic impulses fired by tendon organs. 4. These observations support the assumption that the reduction of Ib autogenetic inhibition in homonymous and synergic motoneurones during GM contractions is due to presynaptic inhibition of transmission in Ib pathways.

Reduction of Ib autogenetic inhibition in motoneurons during contractions of an ankle extensor muscle in the cat.

1. Triceps surae and plantaris (Pl) motoneurons were recorded intracellularly in chloralose or pentobarbital sodium (Nembutal)-anesthetized cats during unfused tetanic contractions of gastrocnemius medialis muscle (GM) produced by stimulating either a cut branch of the GM nerve or the muscle directly. 2. In alpha-motoneurons, during a series of GM twitches at 10/s, contraction-induced inhibitory potentials, probably the result of input from Golgi tendon organs (autogenetic inhibition), rapidly subsided before the end of the series. In contrast, excitatory potentials, probably the result of the activation of spindle primary endings during relaxation from contraction, persisted. 3. In gastrocnemius lateralis-soleus (GL-S) and Pl motoneurons lacking an excitatory connection with Ia afferents from GM, the sustained contraction of this muscle also elicited a declining inhibition. Rapid reduction of contraction-induced autogenetic inhibition was also observed in homonymous gamma-motoneurons. During unfused tetanic contractions lasting 0.5-4s, inhibitory potentials quickly subsided, but an abrupt increase in contractile force elicited a new series of decreasing inhibitory potentials. 4. The assumption that the inhibition induced by GM unfused tetanic contractions was due to activation of homonymous Ib afferents was supported by observations of the effects of electrical stimulation of the GM nerve. In Pl motoneurons lacking an excitatory connection with Ia afferents from GM, repetitive trains applied to the GM nerve, at a strength just above threshold for group I fibers, elicited rapidly declining inhibitory potentials similar to those produced by GM contraction. It was verified that during such stimulation, the amplitude of the group I afferent volleys did not decrease. 5. Reduction of contraction-induced Ib inhibition during sustained GM contraction was still present after a low spinalization of the preparation. As GM tendon organ discharges were verified to persist throughout prolonged contractions, the observed decline of autogenetic inhibition is likely to depend on a spinal mechanism, possibly involving presynaptic inhibition of Ib afferents and/or mutual inhibition of Ib-inhibitory interneurons.

Ensemble discharge from Golgi tendon organs of cat peroneus tertius muscle.

1. The responses of individual tendon organs of the cat peroneus tertius muscle to motor-unit contractions were recorded in anesthetized cats during experiments in which all the Ib-afferent fibers from the muscle had been prepared for recording in dorsal root filaments. This was possible because the cat peroneus tertius only contains a relatively small complement of approximately 10 tendon organs. 2. Motor units of different physiological types were tested for their effects on the whole population of tendon organs in the muscle. Effects of unfused tetanic contractions were tested under both isometric and anisometric conditions. Each motor unit activated at least one tendon organ, and each tendon organ was activated by at least one motor unit. Individual slow-type units were found to act on a single or two receptors, whereas a fast-type unit could activate up to six tendon organs. 3. In one experiment, the effects of 8 motor units on 10 tendon organs were examined. One fast-twitch, fatigue resistant (FR)-type unit acted on six tendon organs, of which four were also activated by another FR unit. The contraction of each unit, on its own, elicited a range of individual responses, from weak to strong. The discharge frequencies of individual responses displayed no clear relation with the strength of contraction, nor did they accurately represent the shape of force profiles. But when all the discharges were pooled, a fairly good correspondence appeared between variations of contractile force and variations of averaged discharge frequencies.(ABSTRACT TRUNCATED AT 250 WORDS)

Postnatal development of peroneal motoneurons in the kitten.

In 1- to 72-day-old kittens, motoneurons of the 3 peroneal muscle nuclei were labeled by retrograde axonal transport of horseradish peroxidase from individual muscles. At birth, the locations of peroneal nuclei were similar to those of the adult cat. Counts of motoneurons at different ages indicated that postnatal cell death does not occur in peroneal motor nuclei. Primary dendrites were as numerous in motoneurons of newborn kittens as in adult motoneurons but they were thinner, shorter and poorly ramified. The number of recurrent axon collaterals was higher in the first postnatal week than at later stages. The growth of motoneurons followed similar rates in the 3 peroneal nuclei. Distributions of cell body diameters and volumes were unimodal at birth and became bimodal between 15 and 20 days postnatal. The separation of peroneal motoneurons in two size subgroups, presumably corresponding to alpha and gamma populations, was followed by an increase in growth rate which became faster for alpha than for gamma motoneurons.

Activation of Golgi tendon organs by asynchronous contractions of motor units in cat leg muscles.

Discharges from individual tendon organs of peroneus tertius muscle activated by the isometric contractions of single motor units were recorded in anaesthetized cats. Pairs of motor units acting on the same tendon organ were stimulated asynchronously at frequencies eliciting unfused contractions. Tendon organ responses to such contractions did not display a linear relation between discharge frequency and contractile force recorded at the muscle tendon. In several instances, one of the motor units exerted a predominant influence on the response of the tendon organ, even though this unit did not produce the strongest activation of the receptor when stimulated on its own.

Unloading of tendon organ discharges by in-series motor units in cat peroneal muscles.

1. The discharges from individual Golgi tendon organs of peroneus tertius and brevis muscles were recorded in anaesthetized cats during the isometric contractions of single motor units. Upon combined contractions of several motor units, two sorts of unloading effects were observed. 2. First, the contraction of a motor unit which, by itself, was without action on a tendon organ could produce a reduction in the response of the receptor to one of its activating motor units. Unloading effects exerted by such in-parallel motor units could effectively interfere with the actions of in-series motor units on the receptor. 3. Second, the contraction of a motor unit activating a tendon organ could reduce the response of this tendon organ to the contraction of another of its activating units. This new type of unloading effect, exerted by in-series motor units, was demonstrated by the fact that the simultaneous contraction of both units elicited less discharge from the receptor than the contraction of a single unit. 4. Unfused contractions of a fast-type motor unit eliciting a response in which the tendon organ discharge was driven 1:1 at the frequency of stimulation of the motor unit, could exert unloading actions on the response of the receptor to another motor unit eliciting a higher discharge frequency. 5. In-series unloading actions were exerted not only by fast-type motor units developing large forces, but also by relatively small slow-type motor units. 6. The high incidence of in-parallel and in-series unloading effects suggests that their consequences may be functionally significant. When large numbers of motor units are being recruited in a muscle, unloading effects might result in a limitation of the Ib afferent discharges from this muscle, preventing an excessive increase of autogenetic inhibition.

Observations on static and dynamic responses of muscle stretch receptors in kittens.

In 5-19-day-old kittens anesthetized with Saffan, the discharges of de-efferented triceps surae muscle receptors were recorded from afferent fibers in dorsal root filaments. The conduction velocities of the afferent fibers ranged between 7 and 30 m/s. Receptors were identified as spindles on the basis of their response to muscle stretch and, whenever possible, the pause in their discharge during muscle contraction. Spindles responding with sustained discharges to muscle extensions of less than 1 mm could be found in 5-8-day-old kittens, provided the muscles was left 'in situ', with intact skin, tendon and aponeuroses. However, such responses were rare before 10 days, and the majority of receptors essentially displayed phasic responses to muscle stretch, in agreement with previous observations. In responses to sinusoidal muscle stretching of 0.1-0.5-mm amplitude, kitten receptors were easily driven to discharge one impulse per cycle at frequencies of 10-15 Hz. But unlike adult spindles, they could not follow higher frequencies unless the amplitude of stretch was increased. The maturation of dynamic responses is slower than that of static responses in kitten spindles. In the first postnatal weeks, small changes in muscle length are poorly signaled to the central nervous system.