Search PubMed⌕ Search

Biomedical subjects

F Clarac

Publications and source records attributed to F Clarac.

At least 73 records · Page 4Linked to original sources

Central control of the sensory afferent terminals from a leg chordotonal organ in crayfish in vitro preparation.

In an in vitro preparation of the crayfish thoracic locomotor system, intracellular recordings have been performed from terminals of a leg joint coxo-basipodite chordotonal organ (CB). In the terminals, some depolarizing events are observed which display all the characteristics of the primary afferent depolarizations (PADs). PADs reduce the amplitude of orthodromic sensory spikes, and thus correspond to a presynaptic inhibition. PADs are tonic in a tonic preparation, and phasic (phase locked) in a rhythmic preparation. A control of the incoming information from the CB could thus be performed by the central nervous system during fictive locomotion.

Action Potentials↗

Fictive locomotion in the fourth thoracic ganglion of the crayfish, Procambarus clarkii.

Bath application of muscarinic agonists induced rhythmic motor activity in an in vitro preparation of the thoracic nervous system of the crayfish, Procambarus clarkii. In 70% of the cases, the rhythm was organized into 1 of the 2 normal patterns: "backward" walking or "forward" walking. In the rest (30%), the ganglion produced either a series of bursts of impulses or no rhythm at all, just an increase in the tonic activity. When it was isolated from all ascending and descending afferents, the fourth thoracic ganglion was still able to generate rhythmic motor output during bath application of muscarinic agonists. In certain motor neurons, muscarinic agonists induced plateau potentials. Under these conditions, some of these motor neurons were able to change the period of the motor pattern, which might suggest that these motor neurons were part of the central pattern generator (CPG) for locomotion. In the presence of 5 x 10(-6)M TTX, the membrane potential of these motor neurons continued to oscillate with organized rhythmic membrane potential oscillations into 1 of the 2 patterns. Under these conditions, current injection into certain motor neurons demonstrated that they continued to affect the CPG. Two classes of walking leg interneurons have been found. First, there are those with a sustained membrane potential: injection of a steady depolarizing current into some of these interneurons induced rhythmic activity in all thoracic motor nerves, even in the absence of any pharmacological activation. Second, there are those with an oscillating membrane potential: these seemed to enable silent motor neurons to be involved in an ongoing rhythm.

Animals↗

Dopamine and motor activity in the lobster Homarus gammarus.

Motor activity similar to agonistic behaviour is obtained after dopamine (DA) injection in lobster. Specially vigorous swimmeret beatings are observed and can be compared to the 'in vitro' motor activity elicited by DA superfusion of the isolated abdominal nervous system. DA-immunoreactive neurons stained by monoclonal antibodies in abdominal ganglia may be involved in swimmeret activation during the agonistic behavior.

Animals↗

Synaptic connections between motor neurons and interneurons in the fourth thoracic ganglion of the crayfish, Procambarus clarkii.

1. A new preparation of the thoracic nervous system of the crayfish, Procambarus clarkii, has been developed, in which it is possible to work with identified members of motor neuronal pools. 2. In such a preparation, it is possible to dissect all specific proximal motor nerves (protractor, retractor, anterior elevator, posterior elevator, and depressor). Motor neurons innervating the four proximal muscles of the fourth walking leg have been identified both physiologically and anatomically by staining the recorded motor neuron with Lucifer yellow through the microelectrode. 3. By the use of cobalt chloride, we have mapped the distribution of somata of all motor neurons within the fourth thoracic ganglion that innervate the different groups of muscles controlling the movement of the fourth walking leg. 4. Most motor neurons innervating the same muscle seem to be electrically coupled, except some depressor motor neurons. 5. Motor neurons innervating antagonist muscles are linked by inhibitory connections. These connections are reciprocal for protractor and retractor motor neurons but usually not reciprocal between elevator and depressor motor neurons. 6. Walking interneurons were identified as neurons without axons in any motor nerve, which modified the motor neuronal activity. Some of them have been injected with Lucifer yellow. 7. Some interneurons make synaptic connections only with antagonist motor neurons that control the movement of one joint. Probably their functional role is to reinforce or to limit the antagonism between each pair of antagonist motor neurons. 8. Other interneurons make synaptic connections with motor neurons innervating muscles controlling different leg joints. These interneurons may play a role in generating the motor patterns that underlie forward and backward walking.

Action Potentials↗

Induction of rhythmic activity in motoneurons of crayfish thoracic ganglia by cholinergic agonists.

In a crayfish thoracic ganglion preparation, it has been possible to induce rhythmic motoneuron (MN) activity by bath application of cholinergic agonists such as oxotremorine or pilocarpine (10(-6) and 10(-5) M respectively). Intracellular recordings of the MN and injection of current pulses demonstrate that these cholinergic agonists are powerful inducers of regenerative properties in MNs some of them being part of the central generator for locomotion.

Animals↗

Single-unit responses and reflex effects of force-sensitive mechanoreceptors of the dactyl of the crab.

This paper examines the responses and reflex effects of force-sensitive mechanoreceptors of the most distal leg segment, the dactyl, of the leg of the crab, Carcinus maenas. The goals of these studies are to establish the potential activities and functions of these receptors in posture and locomotion. The responses of force-sensitive mechanoreceptors to imposed mechanical stimuli depended upon their location on the dactyl. A distal group of receptors is located on a specialized region, the dactyl tip, which is composed solely of epicuticle. Another group of receptors is distributed throughout more proximal regions of the dactyl where the cuticle is completely calcified. Both groups of receptors showed vigorous responses to imposed bending forces. When bending forces were applied as step functions at the dactyl, tip distal receptors showed only phasic responses to all levels of force application. Receptors located at more proximal positions on the dactyl showed phasic responses to low levels of step applied forces and phasicotonic discharges at higher levels of force. Increasing levels of force produced a sigmoid increase in the tonic firing of these units. When bending forces were applied using ramp functions, receptors of the distal group responded with an intense initial discharge followed by firing at a constant rate throughout both force application and release. This response was not related to the velocity of force application. In contrast, receptors located more proximally responded directionally to force application and release. Proximal receptors also effectively encoded the velocity of force application. Responses of these two groups of receptors also differed when vibrations were applied at the dactyl tip: proximal receptors only followed vibrational stimuli up to 50 Hz, whereas distal receptors showed 1:1 responses at vibrations as high as 95 Hz. Mechanoreceptors of the dactyl also responded intensely to bending forces resulting from resisted contractions of the animal's own muscles. No responses were obtained from unresisted movements of the leg. Stimulation of force-sensitive mechanoreceptors of the dactyl produced intra- and interleg reflex discharges in motor neurons to leg muscles. Mechanical bending of the dactyl or electrical stimulation of dactyl nerves in which force-sensitive mechanoreceptors were recorded produced strong tonic excitation of motors neurons to the levator muscles of the same leg.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Force-sensitive mechanoreceptors of the dactyl of the crab: single-unit responses during walking and evaluation of function.

The activities of individual force-sensitive mechanoreceptors of the dactyl (terminal leg segment) of the crab, Carcinus maenas, have been recorded during free walking. These receptors have also been mechanically and electrically stimulated in freely moving animals to directly evaluate their function in locomotion. All force-sensitive mechanoreceptors fired during the stance phase of walking and were silent during swing. Receptor discharges showed regular phase relationships to bursts in motor neurons of leg muscles. Crabs walk laterally and use the legs of one side either in trailing to actively push the animal to the opposite side, or in leading, to less forcefully pull the animal in that direction. Individual force-sensitive mechanoreceptors differed in their patterns of activity during trailing or leading according to their location on the dactyl. Units of proximal receptors fired more vigorously when used in trailing than in leading. Discharges in trailing were also increased by loading of the animal. In contrast, distal receptors near the dactyl tip fired equally intensely during walking in either direction. Proximal receptors thus encode forces and loads applied to the leg. Distal receptors do not encode loads but can signal leg contact and, potentially, exteroceptive vibrations. Sensory stimulation of force-sensitive mechanoreceptors was produced during walking by a device that imposed continuous mechanical bending of the dactyl and by electrical stimulation of dactyl nerves. Intra- and inter-segmental reflexes were evaluated by myographic recordings from leg muscles. Continuous mechanical deformation of the dactyl increased the activity of the levator and decreased firing in the depressor muscles of the homonymous leg during walking. The same stimulus produced enhanced activity in depressor muscles of adjacent legs. The latter effect was not due to simple mechanical coupling resulting from reflexes in the stimulated leg. These reflexes can function to limit forces applied to a leg and provide compensatory adjustments in other legs. Brief low-threshold electrical stimuli applied to nerves in which the activities of force-sensitive mechanoreceptors were recorded produced reflex effects similar to those obtained by mechanical stimulation. These stimuli also reset the rhythm of motor neuron bursting in both homonymous and adjacent legs during walking. These studies confirm the importance of force-sensitive mechanoreceptors in adapting walking patterns and in determining leg coordination in locomotion.

Animals↗

Dual locomotor activity selectively controlled by force- and contact-sensitive mechanoreceptors.

The crab Carcinus maenas walks laterally; moreover, as soon as leg contact with the support is lost, it starts swimming. In free-moving animals, discharges from individual force- and contact-mechanoreceptors located in the terminal segment of the last pair of walking legs have been recorded. These receptors are active during the stance phase of walking and remain silent during swimming. Selective electrical stimulation of their afferent fibers during swimming inhibits this behaviour. The possible role of such sensory information in selecting different motor patterns is discussed.

Animals↗

Single unit sensory activity in free walking crabs: force sensitive mechanoreceptors of the dactyl.

Activities of individual, force-sensitive mechanoreceptors (funnel canal organs) of the terminal segment (dactyl) of the crab leg have been recorded in freely moving animals. During the stance phase of walking receptors discharge in regular bursts that are closely correlated with activity of the opener muscle of the propodite when the leg is used on the trailing side in lateral locomotion. Individual funnel canal organs also show sustained discharges to imposed cuticular strains and strains resulting from resisted muscle contractions. These receptors thus can monitor both internal and external forces that are applied to the leg in locomotion.

Animals↗

Central neuronal projections and neuromuscular organization of the basal region of the shore crab leg.

The musculature and associated skeleton, peripheral nervous system, and central projections of motor and sensory neurones of the two basal (thoracic and coxal) segments of the shore crab leg (fifth pereiopod, P5) were examined in vivo and with methylene blue or cobalt staining. Each of the four main basal muscles, promotor/remotor, levator/depressor, controlling the thoracico-coxal (T-C) and coxo-basal (C-B) limb joints, respectively, comprises several more or less discrete fibre bundles (total 14), with little morphological segregation of different functional groups. The innervation to the basal leg region is carried in two nerve roots arising from the thoracic ganglion. The anterior Th-Cx root carries both sensory and motor axons, while the posterior Th-Cx root is purely motor. Three previously undescribed sensory branches (two "epidermal" nerves and an "accessory" branch), in addition to that innervating the coxobasal chordotonal receptor, have been found in the distal part of the anterior Th-Cx root. Two clusters of 10 to 15 multipolar somata (diam. 30-125 micron) are located proximally at the bifurcation of the accessory nerve and distally where the latter enters the basipodite. The cell bodies (diameter 20-80 micron) of basal leg motoneurones (total ca. 30) lie in the dorsal cortex of the ganglion, with somata of functionally related motoneurones tending to form discrete structural groups. The morphology of individual motoneurones conforms to the general arthropod pattern. All are confined to the ipsilateral hemiganglion and their main neuropilar processes run parallel and in close apposition to each other with overlapping dendritic structures. Sensory projections arising from the CB chordotonal organ also ramify in the region of the neuropile invaded by motoneurones. The possible physiological significance of such structural associations within the CNS is discussed, as are the functional implications of basal limb anatomy in general.

Animals↗

Influence of walking on swimmeret beating in the lobster Homarus gammarus.

Influence of walking on swimmeret beating in intact lobsters, Homarus gammarus, has been analyzed using a treadmill experimental device. Belt movement activates both leg stepping and swimmeret beating. The simultaneity of the onset of the two motor systems in this situation is demonstrated to be the result of a startle response initiated when the belt begins to move. This reaction consists of a non-specific motor activity involving several antagonist postural and dynamic muscles. Abdominal extension and vigorous swimmeret beating are the main features of this reaction. The main characteristics of the swimmeret beating as defined by Davis (1969) has been observed here in sequences without walking. However during long walking sequences a very different swimmeret beating pattern occurs. It is suggested that this slow swimmeret beating is completely subordinate to the walking rhythm during sequences of absolute coordination. In more rapid swimmeret beating a relative coordination with leg stepping is very common. The functional meaning of this linkage between legs and swimmerets is discussed.

Animals↗

Experimental modification of interlimb coordination during locomotion of a Crusacea.

An anlaysis of the phase relationship between the same side legs (homolateral coupling) of Rock Lobster (Crustacea) is performed during sequence of locomtion. On intact animals the phase relationship is similar to the most common insect or mammalian patterns ('alternating model'). After successive autotomy of the walking legs, another coordination pattern occurs progressively; it is an 'in phase model'. From these experiments it can be concluded that there exists a central motor program tightly coordinating each thoracic ganglion and that the alternating pattern could be due to a reorganization of the synchronous system by a peripheral proprioceptive inflow.

Animals↗

Intersegmental reflex coordination by a single joint receptor organ (CB) in rock lobster walking legs.

In the decapod Crustacea, Palinurus vulgaris and Fasus lalandii, the reflex influences of one particular proprioceptor organ, the coxo-basal chordotonal organ (CB), on all the muscles operating the proximal and distal joints of the same leg, have been analysed. The distal end of CB was clamped in fine forceps mounted on a servo-controlled stretcher, and CB length changes of 2 mm were applied. Motor unit activity of the different muscles was recorded as electromyograms (EMGs). 1. Two types of proprioceptive reflex evoked by CB length changes have been investigated: (a) resistance reflexes of the two levator and two depressor muscles of the same leg segment, the coxopodite, i.e. 'intrasegmental reflexes', (b) 'intersegmental reflexes' induced in the muscles operating the proximal (T-C) joint of the same leg, and in all eight muscles of the limb segments distat to CB. 2. Both levator muscles respond reflexly to imposed CB stretch (which normally occurs with limb 'depression'), while both depressors respond during CB shortening (or passive "elevation" of the leg). 3. Intersegmentally CB stretch reflexly activates the M-C extensor muscle, and sometimes facilitates the T-C remotor and C-P bender muscles. Shortening of the single CB organ of a leg excites one or two tonic motor units of the T-C promotor and M-C flexor muscles, and also facilitates the remotor, I-M reductor, and the single stretcher-opener excitatory motoneurone. 4. Some of the muscles, particularly the M-C flexor and extensor muscles, are also influenced intersegmentally by the resting length of CB, usually but not invariably in the same direction as for the corresponding dynamic reflexes. The role of the CB chordotonal organ is discussed, with particular consideration of its intersegmental reflex influence on the posture of the entire leg, and on the more complex motor behaviour of locomotion, where it may be specially significant in coordination of the limb in lateral walking. A complex picture of both tonic and dynamic, inra- and intersegmental reflex regulation of the positions and movements of the limb segments, thus emerges.

Animals↗

Intersegmental reflex actions from a joint sensory organ (CB) to a muscle receptor (MCO) in decapod crustacean limbs.

In the walking legs of decapod crustaceans, intersegmental reflex actions originate from various joint proprioceptors. The activity of the 'accessory flexor' (AF) muscle, which with the myochordotonal organ (MCO) constitutes a muscle proprioceptor for the mero-carpopodite (M-C) joint, is modulated by the sensory discharge of a joint receptor (CB chordotonal organ) for the more proximal, coxo-basal (C-B) joint. Selective mechanical stimulation of the CB organ also reflexly modifies the motor activities of the main M-C flexor and extensor muscles (recorded as EMGs). 1. Dynamic CB stretch (as would occur during a dorso-ventral C-B movement - i.e. 'depression' of the limb) stimulates motor discharge to the M-C extensor muscle, while dynamic release of CB (as during a ventrodorsal C-B movement - or leg 'elevation') excites the accessory flexor as well as the main flexor muscle. 2. Successive M-C muscle responses to repetitive sinusoidal changes of CB length differ quantitatively according to the direction (stretch or release) of the first CB movement, in some cases increasing but more commonly 'adapting' with repetition. 3. Reflex discharge frequencies of the extensor, flexor and accessory flexor motoneurones increase with velocity of CB movement. 4. Eye illumination, and spontaneous or other sources of increased central excitability, generally increase the CB reflex drive to the flexor and accessory flexor muscles and, in parallel, decrease the reflex action on the extensor muscle. The results are discussed in terms of the role of proprioceptive reflexes in intersegmental co-ordination of the leg joints. In particular the significance of the reflex regulation of the myochordotonal receptors, and thereby the gain of the M-C resistance reflexes, is considered in the light of the observed 'co-activation' of main flexor and receptor muscle motoneurones.

Action Potentials↗

[Walking in Crustacea: motor program and peripheral regulation (author's transl)].

1. Rock lobsters can walk in all directions. In the present study, we report the organization of the motor output of the three muscles which control the mero-carpopodite joint (M-C): the extensor E, the flexor F and the accuracy flexor FA, during unrestrained locomotion (fig. 1). 2. During lateral walking, movements of the M-C joint provide most of the propulsive force, whereas during forward and backward walking this joint function more as a strut (fig. 2). Corresponding differences are observed in the motor discharge in the different walking modes. During lateral walking, discharge in the M-C extensor and M-C flexor alternates, whereas during forward and backward walking these antagonists are coactivated (fig. 3 and 4). 3. We have also examined the effects of alterations of proprioceptive feedback: the FA tendon has been cut to eliminate MCO afferents during walking. This ablation does not modify the burst period and the temporal structure of the output pattern is largely unaffected (fig. 5, 6 and 7). MCO may influence the motor output of a given muscle depending upon whether it participates in the return stroke or the power stroke.

Action Potentials↗