Deficits and recovery of body stabilization during acrobatic locomotion after focal lesion to the somatosensory cortex: a kinematic analysis combined with cortical mapping.

We used a kinematic analysis for assessing locomotor impairments and evaluating the time course of recovery after focal injury to the forepaw area of the primary somatosensory cortex (SI) in rats. The animals were trained to traverse a beam that was rotated at various speeds. Changes in orientation of the body and independent movement of the anterior and posterior parts of the body were reconstructed using a 3D motion analysis. In addition, we used electrophysiological cortical mapping to search for neurophysiological changes within the spared cortical zones surrounding the lesion. Neuronal recordings were performed in the same animals prior to and 3 weeks after the lesion induction. Our findings show that a focal lesion that destroyed about 60% of the forepaw representational zone was sufficient to cause conspicuous impairments in the rats' ability to produce adequate motor adjustments to compensate for the lateral shift of the beam and to avoid falling. The main deficits were reflected in a lack of appropriate coordination between the anterior and posterior parts of the body and an inability to maintain a regular gait during locomotion. Skilled locomotion was fully recovered within a 2-3 week period. Functional recovery cannot be ascribed to a restitution of the lost sensory representations. A permanent decrease of forepaw representation was recorded despite the re-emergence of restricted representational sectors in the peri-lesion zone. We suggest that alterations may have occurred in other cortical and subcortical areas interconnected with the injured area. It is also conceivable that the functional recovery involved an increased reliance on all the available sources of sensorimotor regulation as well as the use of behavioral strategies.

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.

Deficits and recovery of head and trunk orientation and stabilization after unilateral vestibular loss.

The aim of the study was to analyse changes in the orientation and stabilization of the head and trunk and their recovery after complete unilateral loss of vestibular information in humans. The ability of nine Ménière's patients to orient and stabilize their heads and trunks in space was investigated during a simple dynamic task of knee-bends and compared with the performance of 10 healthy subjects. Patients' performance was recorded before unilateral vestibular neurotomy (UVN) and during the time-course of recovery (1 week, 1 month, 3 months). Experiments were performed both in eyes open (EO) and eyes closed (EC) conditions to evaluate the role of visual cues in the recovery process. Head and trunk mean angular position (orientation) and mean maximal angular rotation (stabilization) in the roll plane and the yaw plane were recorded using a video motion analysis system. The results indicate that, in the acute stage after UVN (1 week), patients exhibit marked impairments in head and trunk orientation in both visual conditions. In the EC condition, head and trunk were deviated towards the operated side in the roll plane and the yaw plane. Head and trunk stabilization in space was impaired in the roll plane and associated with increased stabilization of the head on the shoulders. Interestingly, vision caused a complete inversion of the orientation pattern, with head and trunk rotations towards the intact side in the roll plane and the yaw plane. Relative to darkness, vision also reduced head and trunk oscillations. Recovery from abnormal head orientation in the light and impaired head stability in both visual conditions was achieved within 1 month and 3 months after UVN, respectively. However, head and trunk orientation in the dark and trunk stabilization in the roll plane remained uncompensated 3 months post-lesion. These results suggest that unilateral vestibular loss leads to a postural syndrome similar to that described previously for various animal species. They confirm the necessity of vestibular inputs for properly stabilizing head and trunk during self-generated displacements in healthy subjects. They also support the notion that vestibular compensation relies on visual cues whose substitution role gradually decreases after UVN.

How changes in vestibular and visual reference frames combine to modify body orientation in space.

The aim of this study was to analyse how changes in vestibular and visual reference frames combine to modify body orientation in space, and to determine the relationship between postural, oculomotor and perceptive parameters. Changes in vestibular and visual references were investigated by comparing controls and vestibular defective patients (Ménière's patients tested before and one week after unilateral vestibular nerve section) under three visual contexts (light with and without vertical and horizontal coordinates, darkness). Unilateral vestibular loss was responsible for postural and perceptive deviations whose direction depended on the presence of visual reference frame. We suggest these changes vary according to the spatial reference frame patients are based on. Postural changes were related to perceptive modifications but not to eye cyclotorsion.

Short-term changes in neck muscle and eye movement responses following unilateral vestibular neurectomy in the cat.

The purpose of this study was to investigate changes in neck muscle and eye movement responses during the early stages of vestibular compensation (first 3 weeks after unilateral vestibular neurectomy, UVN). Electromyographic (EMG) activity from antagonist neck extensor (splenius capitis) and flexor (longus capitis) muscles and eye movements were recorded during sinusoidal visual and/or otolith vertical linear stimulations in the 0.05-1 Hz frequency range (corresponding acceleration range 0.003-1.16 g) in the head-fixed alert cat. Preoperative EMG activity from the splenius and longus capitis muscles showed a pattern of alternate activation of the antagonist neck muscles in all the cats. After UVN, two motor strategies were observed. For three of the seven cats, the temporal activation of the individual neck muscles was the same as that recorded before UVN. For the other four cats, UVN resulted in a pattern of coactivation of the flexor and extensor neck muscles because of a phase change of the splenius capitis. In both subgroups, the response patterns of the antagonist neck muscles were consistent for each cat independently of the experimental conditions, throughout the 3 weeks of testing. Cats displaying alternate activation of antagonist neck muscles showed an enhanced gain of the visually induced neck responses, particularly in the high range of stimulus frequency, and a gain decrease in the otolith-induced neck responses at the lowest frequency (0.25 Hz) only. By contrast, for cats with neck muscle coactivation, the gain of the visually induced neck responses was basically unaffected relative to preoperative values, whereas otolith-induced neck responses were considerably decreased in the whole range of stimulation. As concerns oculomotor responses, results in the two subgroups of cats were similar. The optokinetic responses were not affected by the vestibular lesion. On the contrary, otolith-induced eye responses showed a gain reduction and a phase lead. Deficits and short-term changes after UVN of otolith- and semicircular canal-evoked collic and ocular responses are compared.

What does the haptic modality do during cognitive activities on letter shapes? A study with left- and right-handers.

This study was undertaken to analyze intermanual (interhemispheric) transfer in left and right handed subject and to assess how information was extracted during finger scanning of letter shape at the different levels of letter processing: shape recognition during a physical matching task, letter recognition in a verbal "meaning" matching task and letter naming. The dichhaptic procedure was used to study interhemispheric relations. It was hypothesized that cognitive activities have a feed-forward effect on the exploration of shapes, and that the performance is related to the nature of the task and to handedness. The exploratory strategies of the two types of handedness were also analyzed. The results showed that response latencies were generally similar for left- and right-handed subjects, but accuracy was better for left than right handers in "verbal" matching with the same overall exploratory strategies. In physical matching, left- and right-handed subjects performed equally but used different exploratory strategies. The naming task was very difficult for both groups but failed to discriminate their on accuracy, response latency, and exploratory strategy. The results are discussed with reference to the different exploratory strategies used and the interhemispheric interaction at work in different cognitive processes.

Kinematic analysis of locomotion in unilateral vestibular neurectomized cats.

The vestibular syndrome following unilateral lesion of the vestibular system and the subsequent behavioral compensation over time have been well documented in many species. However, the locomotor pattern changes and the behavioral strategies used to preserve balance have still not been described. This study was aimed at quantitatively describing posturolocomotor behavior in cats tested before and after unilateral vestibular neurectomy (UVN) by the rotating beam test, which provides locomotor tasks of various difficulty. The position of head, neck, and trunk and the trajectory of the forelimbs and hindlimbs were recorded in 5 cats by 3D motion analysis. Step length and frequency walking velocity, and body height were computed. Results showed that normal cats adapted their locomotor patterns to the speed of beam rotation by increasing step length and/or frequency, that is, by increasing walking velocity, but without drastically changing their body posture. By contrast, UVN cats typically lowered their body centers of gravity and modified their locomotor patterns according to the locomotor task. Mean walking velocity was decreased in the low range of beam rotation as a result of smaller step length and lower frequency, and it was increased in the high range by opposite effects on these step cycle parameters. Modifications of the locomotor parameters were a function of the direction of beam rotation, showing significant reduction of step length, frequency, and velocity in the low range of counterclockwise compared to clockwise beam rotation, that is, during rotations toward the lesioned side. Phase plane plots of foot linear velocity with respect to foot linear displacement along the horizontal longitudinal axis displayed two different limit cycles, adapted to easy (low range of beam rotation) and more difficult (high range of rotation) walking conditions, in the normal cat. These dynamic profiles of the trajectories of the limbs during the step cycle were not greatly modified after vestibular lesion, but the phase plane typically observed in the high range for the normals was also found in the low range for the UVN cats. Thus, locomotor equilibrium function in the cat is strongly impaired following UVN, but locomotor balance can still be achieved in the UVN cats by the development of adaptive posturolocomotor strategies compensating for the lack of vestibular inputs.

Alteration of human cutaneous afferent discharges as the result of long-lasting vibration.

The unitary activities of slowly (15 SAI) and fast-adapting type I (12 FAI) skin mechanoreceptive afferent units innervating the anterior part of the human leg and foot were recorded by using the microneurographic method. The recordings were performed both at rest and on application of cutaneous stimuli of various intensities before and after exposure of the corresponding receptive fields to vibration (0.5 mm peak to peak, 100 pulses/s, 10 min). The results show that 11% of the units tested, which were previously silent, developed a bursting pattern of postvibratory activity, which lasted 12 min on average. This induced resting activity may account for the tingling sensations usually perceived after exposure to vibration. Furthermore, application of vibration to the cutaneous receptive fields impaired the response properties of the corresponding cutaneous fibers much more markedly in the case of the SAI than in the FAI units. More specifically, less than one-half of the FAI fibers tested showed a postvibratory depressed sensitivity to skin stroking applied at various velocities that persisted for only a few minutes, whereas the responses of all the SAI units to suprathreshold maintained skin indentations applied with increasing amplitudes decreased significantly for 20 min. These fairly durable changes in the transductive properties of the mechanoreceptive afferent units probably lead to an impairment of perceptual and sensorimotor processes and consequently may at least partly account for the alterations in sensorimotor performance that have been reported to occur in humans after exposure to vibration.

Age effects on reflex and postural responses to propriomuscular inputs generated by tendon vibration.

The effects of aging on two sensorimotor levels of propriomuscular function were investigated in a young (20- to 44-year-old) and an elderly (60- to 86-year-old) population by eliciting segmental reflex and postural responses via the same muscle spindle inflow generated by applying the same pattern of tendon vibration. The latency and amplitude of the reflex responses to vibration (tonic vibration reflex) of the biceps and triceps brachii did not depend on the subjects' age. No major age-related changes were observed in the deep reflexes of the lower limbs. The postural responses to the same vibratory stimulation applied to both the soleus or the tibialis anterior muscles (vibration-induced falling) did not show any changes in latency depending on either age or the visual conditions, whereas the intensity of these responses decreased both with age and when the use of vision was possible. Our results suggest that the two levels at which the same propriomuscular messages were processed are differentially affected by aging. The lower reflex level does not undergo any noticeable impairment, whereas the higher postural control level deteriorates in the elderly, which might be partly responsible for the balance problems which tend to occur more frequently with advancing age.

Visual experience during a sensitive period plays a critical role in vestibular compensation: neuronal substrates within Deiters' nucleus in the alert cat.

In a previous study [31], we showed that Deiters' neurons ipsilateral to a vestibular neurectomy temporarily exhibit increased sensitivity to visual cues about fast movement. It was proposed that this change in the deafferented vestibular neuron response observed only during the first 3 weeks post-lesion plays an important role in the vestibular compensation process. The present study was aimed at analyzing the potential influence over the first 2 weeks post-lesion of visual motion cue deprivation (cats housed in stroboscopic light) and passive visual experience (visual information not correlated to head or body movement) on the visually induced activity of Deiters' cells. The extra-cellular response of single units was recorded during sinusoidal translation of a whole field optokinetic stimulus in six alert cats. Following the deprivation of visual motion cues, vestibular unit responses were found to be tuned to low frequencies of visual stimulation, as in intact cats, and to display a phase lag re. velocity during rapid visual stimulation. Passive visual stimulation was also found to impede the increase in neuronal sensitivity to visual input, although the cats had benefited from normal vision from the 15th day post-lesion. These results are discussed in relation to the functional implication of interactive visual experience within the early stages (sensitive period) of the vestibular compensation process.

Visual sensory substitution in vestibular compensation: neuronal substrates in the alert cat.

The purpose of this study was to investigate adaptive changes in the activity of vestibular nuclei neurons unilaterally deprived of their primary afferent inputs when influenced by visual motion cues. These neuronal changes might account for the established role that vision plays in the compensation for posturo-kinetic deficits after the loss of vestibular inputs. Neuronal recordings were made in alert, non-paralysed cats that had undergone unilateral vestibular nerve sections. The unit responses collected in both Deiters' nuclei were compared to those previously recorded in intact cats. We analysed the extracellular activity of Deiters' nucleus neurons, as well as the optokinetic reflex (OKR) evoked during sinusoidal translation of a whole-field optokinetic stimulus in the vertical plane. In intact cats, we found the unit firing rate closely correlated with the visual surround translation velocity, and the relationship between the discharge rate and the motion frequency was tuned around an optimal frequency. The maximum firing rate modulation was generally below the 0.25 Hz stimulus frequency; unit responses were weak or even absent above 0.25 Hz. From the 4th day to the end of the 3rd week after ipsilateral deafferentation, a majority of cells was found to display maximum discharge modulation during vertical visual stimulation at 0.50 Hz, and even at 0.75 Hz, indicating that the frequency bandwidth of the visually induced responses of deafferented vestibular nuclei neurons had been extended. Consequently, the frequency-dependent attenuation in the sensitivity of vestibular neurons to visual inputs was much less pronounced. After the first 3 weeks post-lesion, the unit response characteristics were very similar to those observed prior to the deafferentation. On the nucleus contralateral to the neurectomy, the maximum modulation of most cells was tuned to the low frequencies of optokinetic stimulation, as also seen prior to the lesion. We found, however, a subgroup of cells displaying well-developed responses above 0.50 Hz. Under all experimental conditions, the neuronal response phase still remained closely correlated with the motion velocity of the vertical sinusoidal visual pattern. We hypothesize that Deiters' neurons deprived of their primary afferents may transiently acquire the ability to code fast head movements on the basis of visual messages, thus compensating, at least partially, for the loss of dynamic vestibular inputs during the early stages of the recovery process.(ABSTRACT TRUNCATED AT 400 WORDS)

Dynamic properties of the vertical otolith neck reflexes in the alert cat.

Electromyographic activity of dorsal neck muscles elicited by sinusoidal vertical linear accelerations was studied in alert cats over a wide range of frequencies. Experiments were performed in head-fixed cats and total darkness in order to activate selectively the otolith system. The polyunitary EMG activity was recorded from splenius capitis muscles in normal and labyrinthectomized cats during vertical translations varying from 0.05-1 Hz with a fixed 290 mm peak-to-peak amplitude. The corresponding accelerations ranged from 0.003-1.2 g. In normal cats, the results showed a bilateral and sinusoidal modulation of the EMG activity characterized by two typical EMG patterns depending on the stimulus frequency. In the low-frequency range (0.05-0.25 Hz), the neck muscles responses were composed of a second harmonic (frequency double that of the input signal: H2 responses). The H2 pattern was characterized by an increase in EMG activity during both the upward and downward parts of translation. These two components of the H2 response were closely related to the two peak velocities (+90 degrees and -90 degrees) of the animal motion. Only slight decreases in amplitude and shifts in phase were observed when increasing the frequency. In the higher frequency range (0.25-1 Hz), the neck muscles response was composed of a fundamental frequency corresponding to the input signal (H1 response). The H1 pattern was in phase with the peak of downward acceleration at 0.25 Hz. A phase lag (up to 45 degrees) and a gain attenuation (16.5 dB) were observed when increasing the frequency. The two H1 and H2 EMG patterns were totally absent in bilateral vestibular neurectomized cats. In unilateral vestibular neurectomized cats, a strong drop in gain and phase advance was noted, which mainly affected the H1 pattern. The present results describe some characteristics of otolith-spinal reflexes acting on the head musculature during vertical motion. They are compared with the neuronal responses that we have recorded within the vestibular nuclei complex in the same experimental conditions. The functional role of the vertical otolith-neck reflexes in stabilizing the head in space during many real-life situations is discussed.

Neuronal coding of linear motion in the vestibular nuclei of the alert cat. I. Response characteristics to vertical otolith stimulation.

The aim of the present study was to investigate some aspects of the central processing of otolith information during linear motion. For this purpose, the response characteristics of 69 vestibular nuclei units to sinusoidal otolith stimulation in the vertical Z axis were analysed in the alert cat. Among this population of neurons which responded to a 0.05 Hz, 290 mm translation, 47 units (70%) displayed a firing rate modulation which followed the input frequency (H1 units). The majority of these neurons exhibited an increase in discharge rate during upward displacement, with a response phase close to the motion velocity or slightly leading downward acceleration. The acceleration related units were divided into two groups according to whether they showed clear increases or only a slight change in discharge rate when the stimulus frequency was increased. The former group was characterized by an average -16.3 dB drop in gain (from 43.9 +/- 1.8 dB, S.D. to 27.6 +/- 7 dB, S.D.) within the 0.05 Hz-0.5 Hz frequency range, while the latter group displayed an average -31.2 dB gain attenuation (from 45.1 +/- 1.1 dB, S.D. to 13.9 +/- 0 dB) within the same decade. In contrast to differences in response gain, all the units tested exhibited a relatively stable phase lead of about 20 degrees with respect to downward peak acceleration. Conversely, units whose response was close to motion velocity in the lower frequency range (0.05 Hz-0.10 Hz) displayed a strong phase lead of about 100 degrees when the stimulus frequency was increased (up to 0.50 Hz). These neurons were thus characterized by an acceleration related response in the higher frequency range. At the same time, an average -24.8 dB gain attenuation (from 47.7 +/- 3.4 dB to 22.9 +/- 3.7 dB) was found in the 0.05 Hz-0.5 Hz decade. The remaining 22 neurons (30%) were called H2 units since they displayed a response waveform double that of the input frequency, a response already described during sinusoidal rotation. Unit discharge reached a peak approximately in phase with maximum upward and downward velocity. Asymmetrical change in unit firing rate about the resting discharge level and different dynamic behavior of the upward and downward response components were usually found. These response characteristics suggest that the H2 patterns are centrally constructed and could result from convergence of otolith afferents having opposite polarization vectors.(ABSTRACT TRUNCATED AT 400 WORDS)

Effects of whole-body vibrations on sensory motor system performance in man.

The effects of whole body vibration (WBV) were studied on subjects trained to perform on tasks involving blindfolded arm positioning (proprioceptive tasks), tracking of visual targets and control of static and dynamic torques. Subjects were vibrated in a seated position by means of a hydraulic jack. The vibration used (0.1 G at floor level and 18 Hz) was that occasionally encountered on medium-size cruising helicopter. The seat was that of a heliccopter pilot whose foam cushion was 6 cm thick with a density of 26 kg/m3. Systematic past-pointing was observed for both arm flexion and extension. Foot and arm visual tracking precision, as determined by position and velocity errors, increased in both directions. Static and dynamic control, rated by torque holding stability and torque amplitude precision, were also significantly altered compared to pre-stimulus readings. The results are interpreted in relation to current knowledge of the effects of vibration induced at spinal, vestibular, and central nervous system levels. It is concluded that the proprioceptive system through which vibration-induced afferents enter the neurological networks is the common denominator for the observed alterations of the position, velocity, and force controls. Our observations suggest that particular care should be taken in helicopters and other vibrating vehicles to prevent vibration from reaching muscular masses, especially those involved in motor tasks.

[Multiunit activity of the inferior olive during a conditioned motor sequence]. / Activité polyunitaire de l'olive inférieure au course d'une séquence motrice conditionnée

Multinuit activity from the inferior olive was recorded in chronic cats during a learned motor task. The animals were trained to perform a succession of rapid flexion-extension arm movements alternating with two maintained postures. No significant differences were observed in the olivary activity during maintained postures. However an increase of activity occurred before the beginning of the flexion detected on the biceps EMG recordings. The first modifications of olivary activity occurred in synchrony with postural reorganization preceding the flexion. This latter involved primarily the triceps. The increase of activity took place during the execution of movement and ended after the reaching of the target.