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Biomedical subjects

L Jami

Publications and source records attributed to L Jami.

At least 19 recordsLinked to original sources

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)

Action Potentials

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.

Afferent Pathways

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.

Afferent Pathways

Alpha and gamma motoneurons in the peroneal nuclei of the cat spinal cord: an ultrastructural study.

The aim of the present study was to investigate whether ultrastructural features can be used as a guide to identify alpha- and gamma-motoneurons among the intermediate-size neurons of the peroneal motor nuclei. The peroneus brevis and peroneus tertius muscles of adult cats were injected with horseradish peroxidase, and motoneurons labeled by retrograde axonal transport were examined by electron microscopy. In both nuclei, the distributions of cell-body diameters, measured in the light microscope, were bimodal covering the range of 28-84 microns, with a trough around 50 microns. The sample of 25 motoneurons selected for the ultrastructural study included not only large (presumed alpha) and small (presumed gamma) neurons but also intermediate-size cell bodies with diameters in the 40-60 microns range. For each motoneuron, 2-5 profiles were reconstructed from ultrathin sections taken at 6-8 microns intervals. Synaptic boutons were counted and their lengths of apposition were measured. On the basis of three criteria, namely: (1) bouton types present on the membrane, (2) percentage of membrane length covered by synapses, and (3) the aspect of the nucleolus, all the examined motoneurons, including those with intermediate sizes, fell into one of two categories. Fourteen motoneurons, with cell-body diameters in a range of 55-84 microns, were contacted by all types of boutons (mainly S-type with spherical vesicles, F-type with flattened vesicles, and C-type with subsynaptic cistern); the synaptic covering of the somatic membrane was over 40% and the nucleus contained a vacuolated nucleolus. These were considered alpha-motoneurons. Eleven motoneurons, with only S and F boutons, a synaptic covering under 30%, a compact nucleolus and a cell-body diameter ranging between 28 and 50 microns, were considered gamma-motoneurons. No other combination of the three criteria was observed. These results show that unequivocal distinction of alpha- and gamma-motoneurons is possible in the peroneal nuclei, on the basis of morphological differences independent of cell-body size.

Animals

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.

Animals

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.

Aging

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)

Animals

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.

Animals

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.

Action Potentials

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.

Action Potentials

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.

Action Potentials

Motor nuclei of peroneal muscles in the cat spinal cord.

The cat peroneal muscles have been used in numerous investigations dealing with the physiological properties of motor units, muscle spindles, and Golgi tendon organs. This report presents a study of the organization of peroneal motor pools in the cat spinal cord by means of retrograde axonal transport of horseradish peroxidase from individual muscles to the corresponding motoneurons. The motor nuclei of peroneus longus (PL), peroneus brevis (PB), and peroneus tertius (PT) muscles formed thin columns in the lateral part of the ventral horn in spinal segments L6-S1. In the transverse plane, the PT and PL nuclei occupied, respectively, dorsolateral and ventromedial positions, with PB nucleus in an intermediate position overlapping with the other two nuclei. Measurements of cell body diameters allowed identification of alpha and gamma subgroups in peroneal motoneuron populations. The average numbers of motoneurons were about 96 alpha and 60 gamma in PL, 75 alpha and 54 gamma in PB, and 34 alpha and 23 gamma in PT. Comparison with data from electrophysiological studies indicated that whole populations of motoneurons were labeled in each motor nucleus. The proportions of gamma motoneurons were the same, and cell bodies of gamma motoneurons had similar sizes in the three peroneal populations. In contrast, alpha motoneurons were significantly smaller in PB than in the two other pools, in keeping with the fact that PB contains a proportion of slow motor units larger than the two other muscles. In large samples of homonymous motoneurons, the numbers of first-order dendrites correlated linearly with motoneuron sizes.

Animals

Further evidence for synaptic actions of muscle spindle secondaries in the middle lumbar segments of the cat spinal cord.

1. The aim of this study has been to investigate the receptor origin of postsynaptic actions evoked by group II muscle afferents in mid-lumbar segments of the cat spinal cord. The experiments tested the hypothesis that the afferents involved were the secondary endings of muscle spindles. 2. Spindle afferents were activated by contractions of intrafusal muscle fibres which were induced by electrical stimulation of fusimotor axons in the distal parts of transected ventral roots by one to three stimuli at 150-500 stimuli/s. A separate series of experiments has shown that such stimuli are effective in activating a considerable proportion of muscle spindle secondaries when contractions of extrafusal muscle fibres are eliminated by differential fatigue of these fibres, provided that several fusimotor axons are stimulated simultaneously. 3. Extracellular field potentials were recorded in the dorsal horn, at such locations where synaptic actions were evoked by electrical stimulation of group II but not group Ia muscle spindle or group Ib tendon organ afferents of pretibial flexors. Effects of activation of spindle afferents following stimulation of fusimotor axons were then compared with effects evoked by electrical stimulation of group II afferents of anterior tibial or extensor digitorum longus nerves and by small stretches of these muscles. 4. Distinct field potentials were evoked by stimulation of ventral root fibres at all locations at which field potentials were obtained from group II afferents stimulated electrically. The latencies of these field potentials were in both cases shorter in the dorsal horn than in the ventral horn. 5. The appearance of these field potentials was not related to contractions of extrafusal muscle fibres and was also observed when these contractions were practically eliminated. Furthermore, their threshold and similar dependence on a potentiating effect of two to three stimuli, as found for single secondaries, allow them to be attributed to secondary endings of muscle spindles.

Action Potentials

Effects of muscle shortening on the responses of cat tendon organs to unfused contractions.

1. The discharges from individual Golgi tendon organs of peroneus tertius and brevis muscles were recorded in anesthetized cats. Responses to unfused isometric contractions of single motor units and combinations of motor units were compared with responses to contractions eliciting muscle shortening (i.e., shortening contractions). 2. In 75% of the examined instances, the effect of muscle shortening during unfused contractions was a slight decrease in tendon organ activation, in keeping with the reduction of contractile tension recorded at the muscle tendon. In other instances there was either no change in tendon organ response or, in less than 10% of instances, a slight increase For two motor units eliciting similar activation of a given tendon organ under isometric conditions, the effect of shortening contraction was not necessarily the same. 3. The reductions observed in tendon organ discharges upon muscle shortening were less than proportional to the reductions of contractile tension and difficult to correlate with the properties of motor units, as determined under isometric conditions. The present observations suggest three main reasons for this lack of relation. 4. The first reason depended on the properties of motor units, in that the relation between length changes and tension changes was not the same for all units. Two motor units developing similar isometric tensions did not necessarily produce the same degree of muscle shortening. Some units produced relatively significant shortening without much loss of tension. 5. Second, the dynamic sensitivity of tendon organs is known to exert a major influence on their responses to isometric unfused contractions, accounting for 1:1 driving of discharge during tension oscillations and high frequency bursts upon abrupt increase of tension. Although less tension was produced and the rate of tension development was slower in shortening contractions, similar manifestations of the dynamic sensitivity of tendon organs were observed. In such cases, the responses of tendon organs were the same whether or not the muscle shortened during contraction. 6. Third, when several motor units were stimulated in combination, the unloading influences of in-parallel units were facilitated by muscle shortening so that unloading effects, which were hardly visible under isometric conditions became evident during shortening contractions.

Animals

[Functional properties of the Golgi tendon organs].

Golgi tendon organs are encapsulated mechanoreceptors present at the myo-tendinous and myo-aponeurotic junctions of mammalian skeletal muscles. Within the tendon organ capsule, the terminal branches of a large diameter afferent fibre, called Ib fibre, are intertwined with collagen bundles in continuity with tendon or aponeurosis at one end. The other end is connected with a fascicle of 5-25 muscle fibres, contributed by several motor units. The contraction of these fibres, exerting strain on the collagenous bundle and causing deformation of sensory terminals, is the adequate stimulus of the tendon organ. For this stimulus, the tendon organ has a very low threshold, so that a single fibre twitch can elicit a discharge from the receptor. A tendon organ can thus signal the contraction of a single one of the 10-15 motor units which contribute fibres to the fascicle connected with the receptor. The number of tendon organs present in a muscle, taken together with the fact that a given motor unit can activate several tendon organs, strongly suggests that the contraction of every motor unit in this muscle is monitored by at least one tendon organ. The exact nature of the information provided by tendon organs to the central nervous system remains an open question because no simple relation could be established between the discharge frequency of a receptor and the contractile forces of its activating motor units. It is known, however, that, due to their dynamic sensitivity, tendon organs are efficient in signaling rapid variations of contractile force. The dynamic parameters of muscle contraction prevail in the information carried by afferent discharges from tendons organs.

Afferent Pathways

[Effects of partial unfused contractions of the gastrocnemius medialis muscle on homonymous and synergist motor neurons in cats].

Autogenetic inhibition of homonymous and synergist motoneurones can be elicited by very weak partial twitches of gastrocnemius medialis muscle, but during sustained contractions the amplitude of inhibitory post-synaptic potentials decreases quickly. A similar decrease also occurs during stronger contractions. The mechanism responsible for this decrease is still active in low spinal preparations. Pre-synaptic inhibition of Ib afferent fibres might contribute to this reduction of efficiency in the transmission of Ib afferent inputs to motoneurones.

Animals

Activation of cat motor units by paired stimuli at short intervals.

1. In adult cats, paired stimulations at short intervals were applied in ventral root filaments to single motor axons innervating the peroneus tertius muscle. Paired impulses were recorded from the muscle nerve simultaneously with the electrical and mechanical responses of the muscle portion of the motor unit (muscle unit). The interstimulus interval was gradually reduced in order to determine the minimum compatible with a full activation of the muscle unit by the second impulse. 2. For motor units of all physiological types, this minimum stimulus interval was the shortest interval allowing initiation and conduction of two impulses in the axon, that is, the absolute refractory period for conduction. Its duration ranged between 0.58 and 0.88 ms, displaying no correlation with the axonal conduction velocity. 3. The amount of tension enhancement produced by paired stimulations at the shortest interval varied with the type of the motor unit: it was largest for fast-fatigable units, intermediate for fast-resistant units and smallest for slow units. 4. Paired impulses elicited by paired stimulations at the shortest possible interval arrived near the muscle at a longer interval because the second impulse was conducted at a slower velocity. The minimum interval between arrival of impulses at the muscle depended on conduction velocity and on conduction distance. 5. In motor axons to peroneus tertius, paired impulses leaving the spinal cord at a mean interval of 0.78 ms arrived near the muscle separated by a mean interval of 1.90 ms. Since such an interval always allowed full activation of the muscle unit by the second impulse, this interval is longer than the refractory period of motor units in this muscle.

Action Potentials