Effect of head position and visual condition on balance control in inverted stance.

This study analyzed the effect of head position and visual condition on the control of balance in handstand, a gymnastics posture that necessitates adaptation of sensory information processing. Five expert gymnasts participated. Centre of pressure trajectories and kinematics of different body segments were recorded. The gymnasts were instructed to maintain three handstands as long as possible in four head positions, with and without vision. Performances and postural stability was much better in the standard and dorsiflexion positions than in the aligned and ventroflexion positions under the two conditions of vision. Performances were lower without vision in the standard and dorsiflexion position. If vision clearly plays an important role, yet the tonic neck reflexes also seem to contribute greatly to control body sways during inverted posture.

Kinematics of treadmill locomotion in rats conceived, born, and reared in a hypergravity field (2 g). Adaptation to 1 g.

The kinematics of treadmill locomotion in rats conceived, born, and raised in a hypergravity environment (HG: 2g) until the age of 3 months was investigated for 5 weeks after their exposition to earth's gravity. The locomotor performance of the HG rats (N=7) was compared to that of age-matched control rats (N=8) housed at 1g for the same period. Kinematic analysis of treadmill locomotion was performed up to 35 days of terrestrial life by an optoelectronic motion analyzer (ELITE system). Results showed that the HG rats exhibited a faster locomotor rhythm (increased number of steps/s), walked closer to the ground, and had a more dorsiflexed foot position. Also, HG rats had shorter steps. The data also highlight a fast adaptation to normal gravity since all the locomotor parameters returned to normal values within 3 weeks. The locomotor modifications may be seen as the persistence of a hypergravity-induced posturo-locomotor adaptation in the centrifuge and/or to more functional changes of sensorimotor systems. Because locomotor performance of HG rats is not severely affected, it is concluded that early development of locomotion processes is highly resistant to gravito-inertial changes.

Behavioural changes induced by early and long-term gravito-inertial force modification in the rat.

The study concerns rats conceived, born and raised in a hypergravity environment (HG: 2 g) for 3 months using a centrifuge. They were then exposed to terrestrial gravity (1 g) and submitted to behavioural tests investigating their spontaneous locomotor activity (open-field), their posture (support surface), and their vestibular function (air-righting reflex). Performances were compared to age-matched control rats housed at 1 g for the same time period. Results showed static and dynamic behavioural deficits as early as the rats were exposed to normal gravity. They exhibited strongly increased motor activity in open-field, with longer travelled distances and more scattered trajectories; in addition, the HG rats displayed more numerous rearings than controls did. They showed postural changes characterized by an enlarged support surface and they did not succeed in the air-righting reflex, due to increased time-delay for head righting. None of these changes were permanent. Indeed, for all tests, the HG rats tested after 3 weeks spent in normal terrestrial gravity exhibited behaviours similar to those of the controls. HG-induced changes in the functional properties of the vestibular system may explain the deficits showed by the HG rats once exposed to normal gravity. The adaptation process to 1 g leading to the appearance of normal behaviour takes about 3 weeks. It likely implicates a central re-evaluation of the sensory inputs and an updating of the motor commands.

Muscular exercise improves knee position sense in humans.

To determine how position sense depends on the functional state of the receptors involved, we assessed the accuracy of knee position sense before and after a moderate exercise on a cycle ergometer. Measurements were done on 32 healthy subjects with four protocols combining two tasks (intramodal: using the contralateral leg, and crossmodal: using a scheme of a leg on a screen) and two ways of positioning (active and passive). Results showed an improved position sense after exercise with the intramodal protocol combined with active positioning of the reference leg. Whatever the mechanisms involved, enhanced motor performances after exercise can be due not only to improved mechanical properties of the muscles but also to better kinesthetic sensibility.

Gait initiation and impairments of ground reaction forces as illustrated in old age by 'La marche à petits pas'.

Ground reaction forces during gait initiation and kinematics of the first step were recorded in 11 elderly patients with idiopathic 'marche à petits pas' and 18 age-matched normal adults. Smaller values of vertical forces, and impaired amplitudes and directions of anteroposterior forces might explain start difficulties of the patients. Higher vertical displacement of the foot, and laborious establishment of the rhythm in the patients could also be related to the perturbations observed in the reaction forces. These perturbations are likely to reflect impairments of muscular synergies of the lower limbs and the lack of limb co-ordination.

Posturo-kinetic effects on kicking movements of a lack of initial ground support under the moving leg.

The posturo-kinetic effects of having no initial ground support under the movement leg were studied during kicking movements. Considerable effects were observed on the efficiency of the voluntary movement, which was noticeably reduced, as well as on the organisation of the postural adjustments. The fact that the efficiency of the movement was lower than under normal conditions shows that the posturo-kinetic reorganisation does not succeed in compensating for the lack of initial thrust of the foot against the floor. These results underline the important contribution of the postural adjustments to the performance of the movement itself, besides their role in maintaining equilibrium and posture.

Relationships between the force of voluntary leg movements and the associated postural adjustments.

Two types of kicking movements were studied: "touch mode" (minimal impact force) and "strike mode" (maximal impact force). Notable differences have been found to exist between the two kicking conditions. In the "strike mode", the onset of the voluntary movement occurred later and the movements of the postural segments preceding the voluntary movement were significantly longer and larger. These results confirm that the early postural adjustments contribute importantly to the efficiency of these movements.

Somatotopical effects of local microinjection of GABAergic agents in Deiters nucleus on the posturokinetic responses to cortical stimulation.

The postural adjustments that accompany the limb flexion following stimulation of the motor cortex in the cat are in part at least mediated by the lateral vestibular nucleus (LVN). In fact, inactivation of vestibulospinal (VS) neurons by unilateral injection of GABA agonists into the LVN decreased the early component of the responses in all of the postural limbs without modifying the threshold, latency, or amplitude of the cortically induced flexion movement. Just the opposite result was obtained after injection into the same structure of GABA antagonists. Experiments were performed to find out whether these VS influences were somatotopically organized. Microinjection of 0.25 microL of the GABA-A agonist muscimol or the GABA-B agonist baclofen (at 1.5 to 4 micrograms/microL saline) into the rostroventral part of the LVN (rvLVN) of one side produced hypotonia in the ipsilateral and hypertonia in the contralateral forelimb. The same injection also reduced the early component of the postural responses to cortical stimulation in both forelimbs, but not in the hindlimbs. On the other hand, unilateral injection of the same agents into the dorsocaudal part of the LVN (dcLVN) produced similar effects, but they were limited to the hindlimbs. The results obtained, which lasted for about 2 to 3 h, were not only site-specific but were also dose-dependent. Injection into the rvLVN or dcLVN of 0.25 microL of the GABA-A antagonist bicuculline or the GABA-B antagonist phaclofen (at 8 or 5 micrograms/microL saline) produced localized changes in posture as well as in the post-urokinetic responses to cortical stimulation. These were opposite in sign to those elicited by the corresponding agonists. In conclusion, it appears that: 1) the motor cortex utilizes descending volleys passing through the LVN in order to elicit the early postural adjustments during a cortically induced limb movement; 2) the resulting VS influences are somatotopically organized; 3) the amplitude of these postural responses can be affected by inhibitory influences probably exerted by Purkinje cells of the cerebellar vermis on the related VS neurons through both GABA-A and GABA-B receptors.

Effect of optic nerve stimulation on neurons in pericruciate cortex of cats.

The distribution of optic chiasm input to different types of neurons in pericruciate cortex of cats agreed with previous work using light flashes. Neuron response times served to differentiate the input pathways to pericruciate cortex, and the types of neurons they influence. Input from the optic chiasm arrived in three distinct surges: the first via the superior colliculus, the second via an unidentified pathway, and the third via the visual cortex. A fourth, diffuse surge arrived in the postcruciate cortex via some unidentified pathway. Stimulation of the contralateral side of the optic chiasm had a weaker effect than stimulation of the ipsilateral side; it evoked activity at a higher threshold, with fewer spikes per response, and at a longer latency. The difference in response latency between the two sides was largest on neurons responding to the first surge, decreasing in later surges, and being least on those neurons responding to the last surge. About 2.3% of the postcruciate and 15% of the precruciate neurons responded only to optic chiasm stimulation; they were isolated in the granular layers, and their responses could not be influenced by prior cutaneous input. It is suggested that much of the visual input to pericruciate cortex serves to modulate on-going cortical output and, thereby, the behavior of the animal.

GABA receptors in Deiters nucleus modulate posturokinetic responses to cortical stimulation in the cat.

The early component of the postural responses which accompany the limb flexion during unilateral stimulation of the motor cortex in the cat is not of reflex origin, but results from a central command. These postural adjustments are characterized by a decreased force under the limb diagonally opposite to the moving one and an increased force under the other two. Since the lateral vestibular nucleus (LVN) exerts an excitatory influence on ipsilateral limb extensor motoneurons, experiments were performed in cats to establish whether the cortical-induced postural changes were mediated through the LVN. This structure is tonically inhibited by GABAergic synapses originating from Purkinje cells of the cerebellar vermis, so that local microinjection into the LVN of GABA agonists or antagonists should either decrease or increase the spontaneous discharge of their neurons. Unilateral microinjection of 0.25 microliters of the GABA-A agonist muscimol or the GABA-B agonist baclofen (at 2-4 micrograms/microliters saline) into the LVN produced a short-lasting episode of ipsilateral postural hypotonia and contralateral hypertonia, during which the cats were unable to stand on the measurement platform. When, shortly after, some recovery of the postural activity appeared, no changes in threshold, latency or amplitude of the cortical-induced flexion movement were observed; however, the early component of the postural responses decreased in the other three limbs. Moreover, the slope of the response curve of the moving limb remained unmodified, while that of the early component of the postural responses, which involved the remaining limbs, decreased following stimulation of the motor cortex at different stimulus intensities. These effects started a few min after the injection and lasted for about 2-3 h. The effects described above were dose-dependent. Moreover, histological controls indicated that the structure responsible for these postural changes corresponded to the middle part of the LVN. The specificity of the results was shown by the fact that unilateral microinjection of 0.25 microliters of the GABA-A antagonist bicuculline or the GABA-B antagonist phaclofen (at 5-8 micrograms/microliter saline) into the LVN produced a postural asymmetry opposite in sign to that elicited in the same experiments by the corresponding agonists. These injections did not modify the amplitude of the cortical-induced limb flexion, but rather enhanced the amplitude of the early component of the postural responses in the other three limbs.(ABSTRACT TRUNCATED AT 400 WORDS)

Injection of a cholinergic agonist in the dorsolateral pontine tegmentum of cats affects the posturokinetic responses to cortical stimulation.

Microinjection into the dorsolateral pontine tegmentum of the muscarinic agonist bethanechol, leading to activation of cholinoceptive pontine reticular formation (pRF) neurons and the related medullary inhibitory reticulospinal system, did not modify the threshold, latency and amplitude of the forelimb flexion elicited by unilateral stimulation of the corresponding motor cortex. However, the remaining limbs which displayed a diagonal pattern of postural adjustments showed a dissociation of their postural responses in 2 components: the early component of central origin greatly decreased in amplitude, while the late component attributed to reflex mechanism triggered by the unbalance brought about by the flexion movement increased. Further evidence indicated that the pRF system intervenes in the gain regulation of the early postural responses during the cortically induced limb movement.

Cholinoceptive pontine reticular structures modify the postural adjustments during the limb movements induced by cortical stimulation.

1. Activation of the pontine reticular formation (pRF) and the related medullary inhibitory reticulospinal (RS) system decreases the postural activity. This effect can be achieved either by local injection into the dorsal pontine tegmentum of cholinergic agonists which excite cholinoceptive pRF neurons, or by injection of noradrenergic agents which block the inhibitory influence exerted by the locus coeruleus (LC) neurons on the pRF. The main aim on the present study was to analyze the effects of tonic activation of these pRF neurons on the postural adjustments accompanying limb movements induced by motor cortex stimulation. In particular, electrodes were implanted chronically in the motor cortex of cats and stainless steel guide tubes of small size, later used for drug injection, were set bilaterally into sites just above the responsive regions. 2. Limb flexion elicited by stimulation of the motor cortex was accompanied by a diagonal pattern of postural adjustment, characterized by a decreased force exerted by the limb diagonally opposite to the moving one and an increased force exerted by the other two. 3. Microinjection into the pRF of both sides of 0.25 microliter of the muscarinic agonist bethanechol at the concentration of 8 or 16 micrograms/microliters in buffered artificial cerebrospinal fluid produced a short-lasting episode of postural atonia followed by a period of reduced postural activity, during which the cats were still able to stand on the measurement platform. Under this condition no changes in threshold, latency and amplitude of the flexion response were observed in the performing limb; however, the postural responses were considerably affected. In particular, when the performing limb was a forelimb, the other anterior limb showed a dissociation of the postural response in two distinct components. The first anticipatory component, which had a short latency (12-15 msec) and was considered to be centrally triggered, decreased in amplitude after injection of bethanechol and sometimes disappeared; on the other hand the second component, which had a long latency (50-60 msec) and was thus considered to be of reflex origin, increased in amplitude, due to the instability resulting from the depression of the early postural response. Similar results also affected to a lesser extent the hindlimbs. Moreover, body oscillations were observed and monitored from the force platforms following the late component of the postural responses.(ABSTRACT TRUNCATED AT 400 WORDS)

Ambulatory computer system for recording and controlling locomotion.

Description of an original miniature computer system capable of recording biomechanical parameters during locomotion, providing auditory signals which can be used in biofeedback locomotor learning, and triggering functional electrical stimulation of nerves and muscles at preselected moments in the locomotor cycle. The system, which is worn on the patient's back, is connected to appropriate gauges with which it is possible to measure knee angle and to record the time of heel contact with the ground. The relevant data are stored in the system memory and can be transferred to a microcassette and later displayed as a function of time on a video screen or printed on paper. The same data can be used on-line to provide a patient with a sound signal as a basis for biofeedback training. This kind of training has been successfully used to correct knee hyperextension during locomotion in hemiplegic patients. The system can also be used to trigger a stimulator and thus constitutes a valuable tool for investigations and treatments involving functional electrical stimulation. Many other applications may be developed, both in basic research and in various fields where motor training is required.

Postural adjustments to head movements in the cat.

Head movements induced by motor cortex stimulation in the cat are accompanied by variations in the vertical force exerted by each limb. These postural responses were found to show stereotyped patterns: with head dorsiflexions an increase was observed in the force exerted by the anterior limbs and a decrease at the posterior limb level. From comparison between the latencies of the force variations, the beginning of head acceleration, and EMG activity in the limb extensor muscles, it was concluded that triggering of these postural responses is not reflex, but depends on the same command as the movement itself. This early response might be a means of avoiding the downward movement of the trunk which would otherwise result from the reaction force corresponding to the upward head movement.

Associated movements, postural adjustments and synergies: some comments about the history and significance of three motor concepts.

Associated movements, postural adjustments and synergies are three motor concepts directly related to each other, although their historical developments have followed relatively independent paths. We show in this study how some notions, often very old, are evolving and become more precise as the number of observations and experimental results increases. We show also how the relationships between these notions and the terms associated with them may vary depending on the historical period and on the authors. The various kinds of muscular activity related to a movement, which are the object of this review, present several aspects, each of them being often described by different terms, which are analyzed. On the other hand we study the various meanings of some terms or expressions, which may correspond to very different aspects of the motor system. The three concepts analyzed have in common a definition from final manifestations (essentially muscular and mechanical) of motor activity. This characteristic is probably due to the lack of enough experimental data to use more refined criteria for definition, like those based upon nervous structures implicated, mechanisms involved or functional significance. The situation may change in the near future, as this field of investigation is now entering a phase of rapid development.

Effects of pressure stimulation of the body surface on posture and vestibulospinal reflexes.

The effects of pressure stimulation of the body surface on postural activities as well as on the response gain of limb extensors to natural stimulation of labyrinth receptors were investigated in intact, as well as in decerebrate cats. In intact, unanesthetized cats, slight pressure applied symmetrically to the body surface at the chest level decreased the tonic activity of the axial (neck) and limb extensor musculature, as well as the proprioceptive reflexes induced by passive flexion of the limbs. The positive supporting reaction caused by pressure applied to the pad of the foot was also depressed. If the cats were suspended in the air by their nape, slight pressure applied to the upper part of the body greatly reduced the tonic contraction of the forelimb extensors to linear acceleration after downward movement of the animal, a response which can be attributed to stimulation of macular receptors located in the sacculus. Moreover, the prominent myotatic reflexes which occurred in all four limbs as soon as the animal touched the floor were greatly depressed, as shown by the fact that the forelimbs displayed only a slight tonic contraction of the extensor musculature during landing, while the hindlimbs collapsed under the weight of the body. In precollicular decerebrate cats there was a good postural activity in all four limbs. Moreover, the multiunit EMG activity of the medial head of the triceps brachii responded to roll tilt of the animal (at 0.15 Hz, +/- 10 degrees) leading to selective stimulation of labyrinth receptors. These responses, characterized by an increased EMG activity during side-down tilt and a decreased activity during side-up tilt, were related to animal position and not to velocity of animal displacement, and are thus attributable to stimulation of macular, utricular receptors. Slight pressure applied to the chest greatly decreased not only the postural activity of the limbs, but also the amplitude of EMG modulation and then the gain in the first harmonic component of the multiunit EMG responses of the triceps brachii to animal tilt. This reduced gain was due, in particular, to a reduced number of motor units being recruited during labyrinth stimulation, although a reduced modulation of firing rate of the active motor units should not be ruled out. However, no changes in the phase angle of the responses were observed.(ABSTRACT TRUNCATED AT 400 WORDS)

Effects of pressure stimulation of the body surface on placing reactions.

The effects of pressure stimulation of the body surface on placing reactions were investigated in intact unanesthetized cats. A slight contact applied to the dorsum of the fore- and hindpaws produced the typical placing reactions in these preparations. These reflexes, however, became more prominent if, in addition to the exteroceptive input, a proprioceptive input was elicited after plantar flexion of the paws. Slight pressure applied to the upper part of the body greatly depressed not only tactile placing reactions, but also to a lesser extent proprioceptive placing reactions. Moreover, these reflexes were less prominent and more sluggish and fatiguable than in the normal cats. The depression of the placing reactions elicited by the cutaneous input during pressure applied to the body surface mainly affected the forelimbs. However, as soon as the pressure stimulation was removed, the placing reactions reappeared or became as prominent as usual. We postulate that the exteroceptive input resulting from body pressure decreased the placing reactions by exciting the Purkinje cells of the intermediate cortex of the cerebellum, which are in turn inhibitory on the interpositus nucleus, thus reducing the neuronal discharge of the interpositorubrospinal pathway acting on flexor motoneurons.

Analysis of vertical displacements of the center of gravity in the cat during limb flexion induced by cortical stimulation.

A limb movement and the associated postural adjustment result in a displacement of the center of gravity of the body. The vertical component of this displacement has been calculated from the variations in the sum of vertical forces at each limb. Through these variations, it is possible to measure the vertical acceleration of the center of gravity. Velocity and displacement are then obtained by two successive integrations of acceleration values. The magnitude of displacement of the center of gravity depends on the magnitude of limb displacement, and hindlimb flexion induces larger displacement than forelimb flexion. It is also directly linked to the time course of the vertical force variations recorded at each limb. The feed-forward character of the postural adjustment appears to minimize this amplitude, which leads us to discuss the functional significance of this postural adjustment.

Changes in posturo-kinetic limb responses to cortical stimulation following unilateral neck deafferentation in the cat.

Unilateral neck deafferentation produces in cats a postural asymmetry, characterized by an increase in the extensor activity of the ipsilateral limbs and a decreased activity in the contralateral limbs; moreover, the placing reactions are severely depressed in the ipsilateral limbs. The present experiments were performed to investigate the influence of unilateral section of the cervical dorsal roots C1-C2 or C1-C3 on the postural adjustments occurring during flexion limb movements induced by cortical stimulation in chronically implanted cats. Only weak stimulations were used and motor activity was mainly isometric, to facilitate quantification of responses analyzed by measuring changes in vertical force exerted at the level of each limb in the up-right position of the animal. The results obtained were the following. The general motor behaviour of the animal was not affected by the lesion. In particular, stimulation of the motor cortex evoked not only flexion movement of a contralateral limb but also an initial postural adjustment consisting of a diagonal force pattern. The response threshold of the moving limb increased on the hypertonic side and decreased on the hypotonic side. Moreover, the changes in threshold of the performing limb following unilateral neck deafferentation were coupled with parallel changes in threshold of the limbs involved in the postural adjustment. There were great quantitative modifications both of the movement and the postural adjustment after the neck deafferentation. In fact weight shift latencies increased when the performing limb was ipsilateral to the hypertonic side, while the weight shifts themselves decreased in amplitude and speed. The opposite occurred when the performing limb was ipsilateral to the hypotonic side. Parallel with these changes, the slope of the response curve of the moving limb as well as of the limbs involved in the postural adjustment decreased following stimulation of the motor cortex at different stimulus intensities if the performing limb was ipsilateral to the hypertonic side, but increased if the performing limb was ipsilateral to the hypotonic side. These changes compensated after chronic neck deafferentation. In summary, the results indicate that well coordinated posturo-kinetic responses were still elicited by cortical stimulation in spite of the prominent changes in posture and reflex movements induced by unilateral neck deafferentation. In particular, changes in threshold, latency, amplitude and speed of the force changes under the limb making a movement were coupled with parallel alterations of the force changes under the limbs undergoing postural adjustment.(ABSTRACT TRUNCATED AT 400 WORDS)