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Biomedical subjects

P Bessou

Publications and source records attributed to P Bessou.

At least 37 records · Page 2Linked to original sources

Discharge patterns of gamma motoneurone populations of extensor and flexor hindlimb muscles during walking in the thalamic cat.

Two monopolar recordings of the whole activity in a fine nerve branch innervating the gastrocnemius lateralis (GL) or the sartorius medialis (SM) muscle were obtained during spontaneous walking in thalamic cats. Using a special electronic device, the potentials of several groups of efferent (alpha and gamma) and afferent (I and II) fibres constituting the whole nerve activity were separated. In the present paper we compare the data obtained for an ankle extensor (GL) and a hip-knee flexor (SM) during the step cycle. In both muscles the gamma motoneurone population is activated in parallel with the alpha motoneurone population. Usually, between the cyclic locomotor discharges, the GL gamma neurones are tonically active whereas the SM gamma neurones are silent. During muscle contraction, the group I and II afferent discharges are both length and gamma dependent, but the prevailing factor is the muscle shortening for the GL afferents and the cyclic gamma drive for the SM afferents. Both dynamic and static fusimotor efferents appear to be activated during muscle contraction, but on indirect evidence it is suggested that dynamic action prevails in GL spindles whereas static action dominates in SM spindles.

Animals↗

Efferent and afferent activity in a gastrocnemius nerve branch during locomotion in the thalamic cat.

The firing patterns of alpha and gamma efferent fibres and of group I and group II afferent fibres innervating the gastrocnemius muscle were 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 whole activity of a thin branch of gastrocnemius lateralis or medialis nerve. The main results were: During the locomotor cycle the activity of the afferent and efferent populations was highly modulated. alpha- and gamma-motoneurones were co-activated within the locomotor cycle during ankle plantar-flexion. The gamma discharge began to rise earlier and to fall later than did the alpha discharge. The amplitude of the gamma discharge, unlike that of the alpha discharge, was largely independent of the vigour of walking. Between the cyclic discharges, most of gamma populations were tonically active whereas alpha populations were silent. Subgroups of the alpha and gamma populations were not usually activated according to the cell-size principle, but, the activation of the latest gamma subgroup always preceded that of the earliest alpha subgroup. Modulation of the group I and II afferent discharges was closely related to the cyclic length changes of the parent muscle. Fusimotor activation during the active shortening of gastrocnemius muscle prevented the afferent discharges from pausing. The pattern of afferent and efferent activity during selective curarisation of the extrafusal junctions indicated that the discharge of static gamma-motoneurones is modulated during the locomotor cycle. After curarisation of both extrafusal and intrafusal junctions, an efferent-discharge pattern of central origin persisted alternately in extensor- and flexor-muscle nerves (fictive locomotion). The durations of the fictive locomotor cycle and of the cyclic discharge in the sartorius nerve were increased as a consequence of the suppression of phasic afferent inputs to the C.N.S. Maintained ankle dorsi-flexion slowed the fictive locomotor rhythm and elicited opposite effects, respectively excitation and depression, on the magnitude of the alpha and gamma discharges. Maintained ankle plantar-flexion scarcely perturbed the duration of the fictive locomotor cycle, but the duration of the sartorius-nerve discharge lengthened at the expense of that of the gastrocnemius discharge. Both gastrocnemius alpha- and gamma-motoneurones were depressed, the former considerably more than the latter. The roles of the gastrocnemius afferents and gamma-efferents during the locomotor cycle are discussed in the light of these results.

Animals↗

Effects of triceps stretch by ankle flexion on intact afferents and efferents of gastrocnemius in the decerebrate cat.

We describe the effects on group I and II afferents as well as alpha- and gamma-efferents of gastrocnemius muscles elicited by stretch and release of the triceps surae muscle performed by a dorsiflexion movement, a maintained dorsiflexion and release of the ankle joint. The experiments were made in decerebrated cats in which the neural loops between the muscles and the spinal cord were intact. Multi-unit discharges of each kind of fibre were obtained, electronically on-line, from two monopolar electrodes 4 mm apart on a small branch of the gastrocnemius lateralis or medialis nerve. Sensory and motor impulses were separated according to the opposite directions of their propagation, then both groups were further subdivided according to conduction velocity. The stretch reflex induced by ankle flexion was associated in eighteen out of nineteen experiments with reflex changes in the activity of the gamma-axon population of the gastrocnemius nerve branch. Facilitatory, depressor and biphasic (facilitatory-depressor) effects were observed, the particular type seeming to depend on the level of gamma tone and gamma-motoneurone size. Flexion release led to cessation of the alpha stretch reflex and to a rebound firing of the gamma-axon population. The lengthening reaction and the clasp-knife phenomenon occurred in certain preparations and were associated with reflex inhibition of the gamma-motoneurones. The over-all responses of group I fibres to triceps stretch showed dynamic and static components, whereas those of the group II fibres were, except in two examples, almost devoid of dynamic sensitivity. During release of stretch and as a result of the simultaneous rebound in gamma activity, the afferent discharges showed a very short pause, no pause, or even a rebound. The origins of the discharges of groups I and II muscle afferents and the factors (muscle length, active and passive muscle tension, reflex changes in fusimotor drive) influencing the activity of the receptors involved are considered as far as possible. Some aspects of the mechanisms of the fusimotor reflex are discussed.

Action Potentials↗

Efferents and afferents in an intact muscle nerve: background activity and effects of sural nerve stimulation in the cat.

1. The background activity was observed in gamma and alpha efferent fibres and in group I and II fibres innervating the muscle gastrocnemius lateralis or medialis. The reflex effects of ipsilateral and contralateral sural nerve stimulations on the muscle efferents were analysed together with their consequences upon the afferents of the same muscle. The observations were made in the decerebrated cat without opening the neural loops between the muscle and the spinal cord.2. The multi-unit discharges of each category of fibres were obtained, on line, by an original electronic device (Joffroy, 1975, 1980) that sorted the action potentials from the whole electrical activity of a small branch of gastrocnemius lateralis or medialis nerve according to the direction and velocity of propagation of the potentials.3. The small nerve may be regarded as a representative sample of different functional groups of fibres conducting faster than 12 m.sec(-1) and supplying gastrocnemius muscles.4. Some gamma efferents were always tonically firing except when a transient flaccid state developed. Usually the alpha efferents were silent, probably because the muscle was fixed close to the minimal physiological length.5. Separate and selective stimulations of Abeta, Adelta and C fibres of ipsilateral and contralateral sural nerve showed that each group could induce the excitation of gamma neurones. The reciprocal inhibition period of alpha efferents during a flexor reflex was only once accompanied by a small decrease in gamma-firing.6. The reflex increase of over-all frequency of gamma efferents resulted from an increased firing rate of tonic gamma neurones and from the recruitment of gamma neurones previously silent. When the gamma efferents in the small nerve naturally occurred in two subgroups, the slower-conducting subgroup (mainly composed of tonic gamma axons) was activated before the faster-conducting subgroup (mostly composed by gamma axons with no background discharge). Some rare exceptions were found, however.7. The selective activation of gamma efferents could be obtained with short-and low-frequency stimulation. When, with stronger stimulations, gamma-alpha co-activation was observed, the onset of the gamma-firing increase preceded by 100-600 msec that of the alpha discharge in the small nerve. Likewise, the onset of the gamma-efferent response preceded the increase of over-all electromyographic activity of the whole triceps muscle but only by 10-100 msec. The discrepancy could be due to the soleus alpha motoneurones being activated earlier than the alpha-motoneurones of gastrocnemius muscle, according to the size principle. In only one experiment, the alpha-firing onset preceded the gamma-firing increase.8. Stimulations of ipsilateral or contralateral nerve, whatever the alpha or gamma reflex patterns, always led to increased firing rates of group I and II fibres of the small nerve. The origins of the discharge of group I and II muscle afferents and the excitation mechanisms of the receptors involved are considered. Some aspects of the mechanism of the reflex control of movement are discussed in the light of these results.

Action Potentials↗

Modification by lidocaine of the discharges of primary endings of muscle spindles in cat tenuissimus muscle.

The effects of lidocaine (1--20 microgram/ml) on afferent discharge patterns of primary endings of muscle spindles in cat tenuissimus muscle were investigated. Discharge from the endings, recorded in Ia afferent axons, was evoked by ramp stretch of the muscle, stimulation of single static or dynamic fusimotor axons or by a combination of stretch and fusimotor stimulation. Spontaneous discharge of the endings at the initial length of the muscles was reduced by 2--5 microgram/ml and abolished by 10--15 microgram/ml lidocaine. The static but not the dynamic discharge elicited by muscle stretch was blocked by concentrations of 10--15 microgram/ml. The same concentrations abolished static and dynamic fusimotor influences on primary ending discharge. However, in one experiment where the spindle was microscopically observed, fusimotor stimulation still resulted in contraction of the intrafusal muscle even when fusimotor stimulation failed to elicit changes in discharge response patterns of the primary endings. These findings indicate that lidocaine interferes with the encoding mechanism prior to block of impulse conduction in either the fusimotor or afferent axons.

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Identification of intrafusal muscle fibres activated by single fusimotor axons and injected with fluorescent dye in cat tenuissimus spindles.

1. Intrafusal muscle fibres of cat tenuissimus spindles have been injected with the fluorescent dye Procion Yellow and identified histologically after recording their changes in membrane potential during 1/sec stimulation of single static or dynamic gamma axons. 2. Thirteen intrafusal muscle fibres innervated by static gamma axons were identified as eight bag2 and five chain fibres. The fact that none proved to be a bag1 fibre is not regarded as significant, for reasons given in the Discussion. 3. In one spindle Procion Yellow was injected into two intrafusal muscle fibres activated by the same static gamma axon; they were identified as a bag2 and a chain fibre. 4. Nine intrafusal muscle fibres innervated by dynamic gamma axons were identified as seven bag1 fibres, one bag2 fibre, and one long chain fibre. 5. In one spindle two bag fibres were injected, one activated by a dynamic gamma axon, the other by a static gamma axon; the former proved to be a bag1 fibre, the latter a bag2 fibre. 6. Stimulation of static gamma axons elicited junctional potentials in seven bag2 fibres and one damaged chain fibre, and action potentials in one bag2 and four chain fibres. In the whole sample of impaled intrafusal muscle fibres (identified and unidentified) activated by static axons, junctional potentials were recorded from twenty-three (62.2%), and action potentials from fourteen (37.8%). Stimulation of dynamic gamma axons always elicited junctional potentials. 7. In a number of instances it was possible to examine the ultrastructure of motor endings belonging to the stimulated gamma axon. The myoneural junctions of trail endings supplied by static gamma axons to bag2 and chain fibres were both smooth and folded; the deepest and most regular folding occurred on chain fibres. The terminals of p2 plates supplied to bag1 fibres by dynamic gamma axons had smooth myoneural junctions.

Action Potentials↗

Histological analysis of cat muscle spindles following direct observation of the effects of stimulating dynamic and static motor axons.

1. Eleven cat tenuissimus spindles have been analysed mainly by cutting serial, transverse, 1 micrometer thick sections following direct observation of the effects of dynamic motor (gamma or beta) stimulation. 2. Histological results from these spindles were also used to interpret the effects of static fusimotor stimulation of other spindles. 3. Dynamic motor stimulation usually produced contractions seen as convergent movements of sarcomeres in single bag fibres, identified as bag1 fibres for reasons given in the text. 4. In one spindle a single dynamic axon produced a translational movement in one pole of a bag1 fibre and a convergent movement in each pole of a bag2 fibre, together with movements in other unidentified (presumably chain) fibres. Subsequent analysis showed that besides innervating both bag fibres the axon also supplied two chain fibres. 5. Contrary to expectation, motor endings on the bag1 fibres seldom occurred at the sites of convergent movement. Only two cases of coincidence occurred among sixteen foci and twenty-one motor endings; otherwise focus and nearest ending were separated by distances of 0.85--2.5 mm. 6. Most of the convergent movements of sarcomeres observed in bag1 fibres occurred in a region of the pole that is ultrastructurally distinct from the region where most of the motor endings were located. The possible relevance of this to the production of contractions in the bag1 fibre is discussed. 7. Convergent movement foci in bag2 fibres produced by the stimulation of static axons occurred largely within the same regions of the pole as the motor endings were located, though, whereas foci were observed in both intra- and extracapsular regions, most of the endings were intracapsular.

Animals↗

Cinematographic analysis of contractile events produced in intrafusal muscle fibres by stimulation of static and dynamic fusimotor axons.

1. Muscle spindles with an intact blood supply and uninterrupted connexions with ventral and dorsal spinal roots (Bessou & Pagés, 1967, 1972) have been prepared in cat's tenuissimus muscles with the aim of cinephotographically recording intrafusal movements induced by the stimulation of single static or dynamic gamma axons; the time cours of these movements and the morphological kind of activated intrafusal muscle fibres have been established. 2. Displacements of spindle guiding marks in the equatorial region elicited by stimulating single static gamma axons are 4-20 times greater in amplitude than the ones elicited by stimulating dynamic gamma axons at the same frequency. 3. The dynamic gamma axons induced a contraction only in nuclear bag fibres which, in addition, never received any static gamma innervation. The static gamma axons evoked contractions either in nuclear bag fibres alone, or in nuclear chain fibres alone, or in both types of intrafusal fibres. Two thirds of static gamma axons supplied nuclear bag fibres. For various reasons, one half only of static gamma axons innervating nuclear bag fibres could be shown to simultaneously innervate nuclear chain fibres. Consequently, about one third of static gamma axons supplied both nuclear bag fibres and nuclear chain fibres, but it is highly probable that this latter figure is an underestimate. One third of static gamma axons produced contraction in nuclear chain fibres only. In this work, the distribution of fusimotor axons has been established in only one muscle spindle of the cluster of muscle spindles that each fusimotor axon is generally innervating. 4. Generally speaking, a static gamma axon elicits contraction of several intrafusal fibres whereas a dynamic gamma axon innervates only one intrafusal fibre and frequently only one pole of the fibre. 5. One third of static gamma axons evoked contractions in nuclear chain fibres that seemed to involve the whole pole. The other static gamma axons and all dynamic gamma axons produced, in the intrafusal fibres that they supplied, one or several foci of localized contractions. 6. The nuclear chain fibres contract and relax faster than nuclear bag fibres. The contractions of nuclear bag fibres supplied by static gamma axons are stronger and faster than those of nuclear bag fibres innervated by dynamic gamma axons. Nearly all nuclear bag fibres innervated by static gamma axons, like the nuclear chain fibres, show transient contractions at each pulse of a stimulation at low frequency (2-20/sec). 7. The results are discussed taking into account the available anatomical and physiological data on the muscle spindle. Their consequences with regard to intrafusal working are briefly considered.

Animals↗

Intracellular potentials from intrafusal muscle fibers evoked by stimulation of static and dynamic fusimotor axons in the cat.

1. Membrane potential changes of single intrafusal muscle fibres were intracellularly recorded in spindles of cat's tenuissimus muscle on stimulating single static or dynamic fusimotor axons.2. One third of responses elicited on stimulating static fusimotor axons were action potentials while the remainder were junction potentials. Repetitive stimulation of static axons eliciting junction potentials produced summation, facilitation and in some instances the appearance of propagated potentials.3. All the responses evoked on stimulating dynamic fusimotor axons were junction potentials which summated but never produced propagated potentials during repetitive stimulation.4. Most of the impalements leading to junction potentials were located in the transition zone between intra- and extra-capsular regions of spindle poles.5. The relation between extracellular and intracellular potentials elicited by stimulation of a fusimotor axon (static or dynamic) makes it possible to assign a physiological nature to the junction potentials recorded intracellularly and to exclude effects attributable to injury.6. A coupling between junction potential and local contraction is indirectly inferred from the frequencygrams which have been previously obtained during single shock stimulation of dynamic fusimotor axons.

Action Potentials↗