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Biomedical subjects

A Berthoz

Publications and source records attributed to A Berthoz.

At least 19 recordsLinked to original sources

Cortical control of vestibular-guided saccades in man.

Memory-guided saccades, made to a remembered location to which gaze was directed before a passive body rotation (i.e. with a vestibular input), were electro-oculographically recorded in 24 patients with various cortical lesions and in 18 control subjects. Anticipation and latency, direction errors and accuracy of the first saccade, stability of eye position in darkness and final eye position were quantified. Patients were divided into small groups, each with lesions affecting one of the following cortical areas: left or right frontal eye field (FEF), left or right prefrontal cortex (area 46 of Brodmann) (PFC), left supplementary eye field (SEF), left or right posterior parietal cortex (PPC) and right parieto-temporal cortex (PTC). There were some abnormalities in the results of the right FEF group, concerning anticipation, direction errors and latency of the first saccade, but no abnormality in the accuracy of the first saccade or of the final eye position. Results in the left FEF group were normal. Accuracy of the first saccade was impaired in the SEF group, bilaterally. Final eye position was also inaccurate in the SEF group. In both PFC groups, significant and, in general, bilateral abnormalities existed for all tested parameters. Accuracy of the first saccade was impaired in the PTC group, leftwards. In contrast, the results in both PPC groups were not significantly different from those of control subjects. Our results suggest that (i) the PFC is involved in the memorization of saccade goals probably encoded in spatiotopic coordinates; (ii) the SEF, but not the FEF, is involved in the control of accuracy of these vestibular-derived goal-directed saccades; (iii) the PTC (i.e. the vestibular cortex), but not the PPC, is involved in the control of such saccades. Therefore, a cortical network different from that involved in the control of memory-guided saccades made to visual targets, with only the PFC in common, could control vestibular-derived goal-directed saccades.

Adult

Effect of post-training unilateral labyrinthectomy in a spatial orientation task by guinea pigs.

The effects of unilateral labyrinthectomy in guinea pigs have been studied on an angular orientation task consisting, in an open field, of running to a hidden goal oriented at 45 degrees with respect to the cephalocaudal axis of the animal placed in a starting-box. The task was conducted in light but in an homogeneous environment, i.e. without visual, auditory or olfactory cues indicating the location of the goal. A second group of animals was submitted to a similar task running to a hidden goal but the place of the goal was indicated by a colored card. All the animals were trained before the lesion and tested in their respective task for 1 month after the lesion. In the task conducted without conspicuous cues, animals were dramatically disturbed. In contrast, animals pretrained in the visually guided task were not impaired after the lesion. These results point out the important role of vestibular information in performing spatial tasks based on angular estimation, since, even if proprioceptive and visuokinesthetic information remain available, subjects seemed not able to maintain a correct angular trajectory. The trajectories being not disturbed in the visually guided task, one can exclude the hypothesis that such deficit was due to a purely motor disturbance.

Animals

A neural network model of sensoritopic maps with predictive short-term memory properties.

Coordinated orienting movements can be accurately performed without direct sensory control. Ocular saccades, for instance, have been shown to be reprogrammed after target disappearance when an intervening eye movement is electrically triggered before the saccade onset. Saccadic eye movements can also be executed toward memorized targets, even when the subject has been passively moved in darkness. Two hypotheses have been proposed to account for this goal-invariance property: either (i) the goal is reconstructed and memorized in the stable frame of reference linked to the environment ("allocentric, coordinates") or (ii) the goal is selected and memorized in the sensors-related maps ("egocentric coordinates") and is continuously updated by efferent copies of the motor commands. In this paper, we shall describe a formal neural network based on this second hypothesis. The results of the simulation show that target position can be memorized and accurately updated in a topologically ordered map, using a velocity-signal feedback. Moreover, this network has been submitted to a simple learning procedure by using the intermittent visual recurring afferent signal as the teaching signal. A similar mechanism could be involved in control of limb movement.

Computer Simulation

Eye and head coupled and dissociated movements during orientation to a double step visual target displacement.

Tight coupling between eye and head movements has been observed in response to a single visual target offset. On this basis, when the visual stimulus consists of two successive steps in the same (horizontal) direction, either increasing in eccentricity (staircase) or decreasing in eccentricity (pulse-step) gaze should be due to concomitant eye and head angular displacement. That is, the eyes and head should aim at each target displacement so that their combined movement matches target offset. We have tested this hypothesis in five healthy subjects. The measured variables were head and gaze offset, the interval between two consecutive saccades from onset to onset (I) and the response delay between onset of the second step and onset of the first gaze saccade (D). With both staircase and pulse-step stimuli, the eye saccade preceded the head movement, and the gaze response either had the stimulus profile pattern or consisted of one gaze saccade to the final target offset. In response to staircase stimuli, I decreased concomitantly with an increase in D; with pulse-step stimuli, as D increased, I decreased slightly in three subjects and decreased markedly in two subjects. Dissociation between the eye and head movements could clearly be demonstrated with pulse-step stimuli: the first gaze saccade to the target pulse displacement was accompanied by a head movement to the target step offset. We also observed cases in which the gaze saccade to the target step displacement was made simultaneously with the head movement to the target pulse offset. Our study extends previous observations in head fixed condition and illustrates that in the majority of cases, when the head is free and a visual pulse step stimulus is presented, both the saccadic and head systems have the ability to modify or cancel the initial neural command to move to the first target displacement. When this modification takes place in only one system, eye and head movements are dissociated.

Electrooculography

Head stabilization during various locomotor tasks in humans. II. Patients with bilateral peripheral vestibular deficits.

This experiment, which extends a previous investigation (Pozzo et al. 1990), was undertaken to examine how head position is controlled during natural locomotor tasks in both normal subjects (N) and patients with bilateral vestibular deficits (V). 10 normals and 7 patients were asked to perform 4 locomotor tasks: free walking (W), walking in place (WIP), running in place (R) and hopping (H). Head and body movements were recorded with a video system which allowed a computed 3 dimensional reconstruction of selected points in the sagittal plane. In order to determine the respective contribution of visual and vestibular cues in the control of head angular position, the 2 groups of subjects were tested in the light and in darkness. In darkness, the amplitude and velocity of head rotation decreased for N subjects; these parameters increased for V subjects, especially during R and H. In darkness, compared to the light condition, the mean position of a line placed on the Frankfort plane (about 20-30 degrees below the horizontal semi-circular canal plane) was tilted downward in all conditions of movement, except during H, for N subjects. In contrast, this flexion of the head was not systematic in V subjects: the Frankfort plane could be located above or below earth horizontal. In V subjects, head rotation was not found to be compensatory for head translation and the power spectrum analysis shows that head angular displacements in the sagittal plane contain mainly low frequencies (about 0.3-0.8 Hz). The respective contribution of visual and vestibular cues in the control of the orientation and the stabilization of the head in space is discussed.

Adult

Eye-head coupling in humans. II. Phasic components.

A tonic coupling between the horizontal component of eye position and dorsal neck muscle activity has been demonstrated in animals and humans. In addition, a transient saccade related coupling has been found in animals. In order to investigate such a phasic component of the eye-head synergy in humans, we have recorded the activity of isolated motor units in the splenius muscle during large horizontal eye movements in head fixed subjects. Eye movement recording was achieved by conventional binocular electro-oculography and the activity of the right splenius muscle was recorded with Bronks coaxial electrodes inserted manually at the C4-C5 intervertebral level. We found two main types of motor unit discharge patterns in the splenius (SPMU), the first type (type A, 14 SPMUs) shows a phasic modulation of firing rate during saccades with a triphasic profile composed of a pre-saccadic suppression, a per-saccadic burst and a post saccadic tonic discharge proportional to eye position. The second type (type B, 6 SPMUs) exhibits little, if any, modulation of firing rate with either fixation or saccades. These results suggest that eye-head coupling is present not only during the fixation period but also during saccades and that a phasic activity or suppression related to saccadic eye velocity is present in dorsal neck muscle EMG.

Adult

Head stabilization during locomotion. Perturbations induced by vestibular disorders.

Head kinematics was studied in 10 normal subjects (NS) and 7 patients (P) with bilateral vestibular deficit while they executed various locomotor tasks. The movement of the body was recorded with a video system which allowed a computer reconstruction of the motion of joint articulations and other selected points on the body in three dimensions. Analyses focus on head translation along the vertical axis and rotation in the sagittal plane. Two conditions were studied: free walking (W) and hopping (H). The subjects were tested in light and in darkness. In NS, while walking in darkness, mean head position was tilted downward. In contrast, this flexion was not systematic in P. Darkness did not significantly influence the amplitude and velocity of head angular displacement during W, but, during H the amplitude decreased by 37% for NS. During H in darkness, head stabilization decreased for P. These results suggest that head kinematics, during natural locomotor tasks, could be used to evaluate vestibular deficiencies.

Adult

Head stabilization during various locomotor tasks in humans. I. Normal subjects.

Head kinematics were studied in ten normal subjects while they executed various locomotor tasks. The movement of the body was recorded with a video system which allowed a computer reconstruction of motion of joint articulations and other selected points on the body in three dimensions. Analyses focus on head translation along the vertical axis and rotation in the sagittal plane. This was done by recording the displacement of a line approximating the plane of horizontal semi-circular canals (the Frankfort plane: F-P). Four conditions were studied: free walking (W) walking in place (WIP) running in place (R) and hopping (H). In the 4 experimental conditions, amplitude and velocity of head translation along the vertical axis ranged from 1 cm to 25 cm and 0.15 m/s to 1.8 m/s. In spite of the disparities in the tasks regarding the magnitude of dynamic components, we found a significant stabilization of the F-P around the earth horizontal. Maximum amplitude of F-P rotation did not exceed 20 degrees in the 4 situations. Vertical angular velocities increased from locomotion tasks to the dynamic equilibrium task although the maximum values remained less than 140 degrees/s. Predominant frequencies of translations and rotations in all the tasks were within the range 0.4-3.5 Hz and harmonics were present up to 6-8 Hz. During walking in darkness, mean head position is tilted downward, with the F-P always below the earth horizontal. Darkness did not significantly influence the amplitude and velocity of head angular displacement during W, WIP and R, but during H the amplitude decreased by 37%. Residual head angular displacement is found to compensate for head translation during the 4 conditions. Our study emphasizes the importance of head stabilization as part of the postural control system and described as a basis for inertial guidance.

Adult

A neurophysiological study of prepositus hypoglossi neurons projecting to oculomotor and preoculomotor nuclei in the alert cat.

The activity of 62 antidromically identified prepositus hypoglossi neurons was recorded in 10 alert cats during spontaneous, vestibular or visually induced eye movements. Neurons were antidromically activated from stimulating electrodes implanted in the ipsilateral medial longitudinal fasciculus (n = 24), the ipsilateral interstitial nucleus of Cajal (n = 6), the ipsilateral parabigeminal nucleus (n = 2), the contralateral superior colliculus (n = 6) and the contralateral cerebellar posterior peduncle (n = 24). Neurons were identified as eye-movement-related when their rate-position and/or rate-velocity plots showed correlation coefficients greater than or equal to 0.6. They were further classified as "position", "position-velocity" and "velocity-position" according to their relative eye position and velocity coefficients. However, they seemed to be distributed as a continuum in which a progressive decrease of eye velocity sensitivity was accompanied by a proportional increase in eye position sensitivity. "Position-velocity" neurons (n = 9) were mainly horizontal type II neurons projecting to the vicinity of the oculomotor complex; two of these neurons with vertical sensitivity were also activated from the interstitial nucleus of Cajal. Mean position and velocity sensitivity of these neurons were 5.2 spikes/s per degree and 0.62 spikes/s per degree per second, respectively. Pure "position" neurons (n = 7) also showed activation during ipsilateral eye fixations; their mean position gain was 7.3 spikes/s per degree and they projected to the ipsilateral oculomotor and Cajal nuclei, and to the contralateral superior colliculus. "Velocity-position" neurons (n = 18) were type I or II neurons with rather irregular tonic firing rates and a mean velocity gain of 0.75 spikes/s per degree per second. Type II "velocity-position" neurons projected mainly to the oculomotor area, while type I neurons projected preferentially to the cerebellum. A special type of "pause" neuron (n = 5), with very low firing rate and pausing mainly for contralateral saccades, was activated exclusively from the contralateral posterior peduncle. Many neurons with weak eye movement sensitivity (n = 22) were activated mainly (73%) from the cerebellum. It can be concluded that the prepositus hyperglossi nucleus distributes specific eye movement related signals to motor and premotor brainstem and cerebellar structures. The variability of interspike intervals of representative prepositus hypoglossi neurons of each class was compared to the discharge variability of identified abducens motoneurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Abducens Nerve

Head kinematic during various motor tasks in humans.

Head kinematic during various motor tasks was studied in ten subjects. The movement of the body was recorded with a video system (E.L.I.T.E.) which allows a computer reconstruction of three-dimensional motion of selected points on the body. Analysis is focused on head rotation in the horizontal and vertical planes. The results demonstrate that the amplitude and the maximum velocity do not exceed respectively 38 deg/s and 185 deg/s. However the head is intermittently stabilized and the angle of this stabilization is dependent upon the task and related to the direction of gaze. Darkness had no significant effect on head rotational velocity during walking but caused a decrease in velocity during running and hopping. The results suggest that head stabilization (1) is related to an ocular fixation point in the direction of gaze in space and (2) is probably regulated on the basis of a predictive mode of sensory motor control.

Adult

Neural correlates of horizontal vestibulo-ocular reflex cancellation during rapid eye movements in the cat.

1. The aim of the present study is to describe the behaviour of identified second-order vestibular neurones in the alert cat during eye saccades. A selection of neurones which are involved in horizontal eye movements has been made. The activity has been compared with a selected sample of abducens motoneurones recorded in the same animals. 2. Alert head-fixed cats were used for this study. Eye movements were recorded by the scleral search coil technique. Abducens motoneurones were identified by antidromic stimulation from the VIth nerve with chronically implanted electrodes. They were recorded extracellularly. 3. Second-order vestibular neurones were identified by orthodromic stimulation from the vestibular organs. They were recorded intra-axonally and injected with horseradish peroxidase after recording of their physiological characteristics. Their morphology was reconstructed from frozen sections. 4. All the recorded vestibular neurones showed various amounts of eye position sensitivity. The firing rate (F) - horizontal eye position (H) characteristics are compared for abducens and vestibular neurones. The population average values are F = 33 + 4 H for motoneurones and F = 51 + 2.4 H for vestibular neurones. 5. All recorded vestibular neurones showed an increase of discharge rate during contralateral horizontal saccades and a strong decrease or pause during ipsilateral saccades. Firing rate - horizontal eye velocity sensitivity has been calculated. 6. Results suggest a strong inhibitory input on vestibular neurones from the saccadic generator. This mechanism underlies the suppression of the vestibulo-ocular reflex during saccades. Our results suggest that in the cat, for saccades of amplitude smaller than 20 deg, there is a variable degree of suppression which is provided by a projection of excitatory bursters (EBNs) on second-order vestibular neurones through inhibitory type II neurones. 7. We also conclude from this study that the eye position sensitivity of vestibular second-order neurones is in fact a motor signal indicating a motor error, i.e. the amount of head or eye movement which remains to be done in order to align gaze on target with the eyes centred in the orbit.

Abducens Nerve

Contribution of the otoliths to the calculation of linear displacement.

1. The present work is a quantitative study of the eye movements induced by linear translation when the subject is instructed to stabilize his gaze on a memorized earth-fixed target. These experiments may allow a better understanding of the central processing of otolithic signals. 2. Human subjects were submitted to either sinusoidal or step-like horizontal linear displacements along the interaural (Y)-axis in darkness, seated in a cart moving along a linear track. Each subject's head was fixed by a helmet secured to the cart. They were asked to keep their eyes on an earth-fixed memorized target at 63 cm from them on the X-axis. 3. During sinusoidal motion, a combination of low smooth compensatory eye movements and of compensatory saccades allowed the subjects to track the memorized target. The linear model of the responses of five subjects (seven sessions) exhibited a near-ideal slope of 1.14 (range 0.84-1.58). Two subjects did not compensate properly for their displacement. The mean "vestibular-saccadic" (VS) gain (ratio of overall eye movement peak-peak amplitude versus head displacement amplitude) was 1.52 +/- 0.80 (SD), showing an overestimation of head displacement. 4. The otolith-ocular reflex (OOR) mean gain values (ratio of slow phase cumulated peak-peak amplitude versus head displacement amplitude) were about 0.13 degrees/cm. This value is 5 times higher than what has been reported in the literature, probably due to the fact that the target was at a short distance. 5. The number of saccades occurring during sinusoidal stimulations varied according to the different subjects. They were obviously compensatory saccades and not quick phases. They indicate that although the gain of the OOR was small, the brain has computed the adequate desired eye position. 6. During steplike head displacements in darkness, although the OOR gain was also small, seven of the eight subjects could stabilize their gaze with a mean VS gain of 1.01 +/- 0.70. The linear model for the pooled responses of these subjects exhibited a slope of 0.82. 7. When subjects were instructed not to move their eyes during the translation, three of the five examined could still correctly reproduce the head movement amplitude with saccades, even as late as 50 s after motion had stopped. This indicates that head displacement was stored with the adequate metrics and could be used to drive the saccadic system. 8. Bilabyrinthectomized patients could not perform any adequate gaze stabilization. This shows that the observed performance was of vestibular origin.

Adult

[Cooperation and substitution of the saccadic system and the reflexes of vestibular origin: should the "reflex" concept be revised?].

Neuronal mechanisms of production of ocular saccadic movements are discussed, together with stabilizing movements of vestibular origin, and descriptions given of several neuronal elements constituting the tecto-reticulo-spinal system, the generator of horizontal saccadic movements in the brain stem, the vestibulo-ocular pathway, etc... Analysis of these pathways and of the signals travelling through them suggests that the overall apparatus is not constituted of isolated reflex modules but acts as a true functional unit for cooperation between orientation and stabilization mechanisms. In addition, coordination of synergies between eye and head movements appear to be based on the activity of clearly identifiable neurons. This neuronal implementation of the concept of synergy is completed further by a remarkable functional flexibility. The latter allows substitution by the brain, for example, fo saccades instead of vestibular reflex movements when the latter are deficient because of lesions or disturbances in the coherence of sensory information. The principal conclusion reached is that cognitive factors, which imply representations of body movements in space activated as a function of the subject's goals, must be considered when ocular movement, and gaze control is described.

Animals

Changing patterns of eye-head coordination during 6 h of optically reversed vision.

1) This study investigates the early development of adaptive changes in oculomotor function associated with coordinated eye-head tracking of the optically reversed image of an earth-fixed target seen through horizontally reversing dove prism goggles attached to the skull. 2) Two tasks comprised a) fixation of a single target during head rotation which causes the seen target's image to move in the direction of head motion by an amount exactly equal to the head movement itself (the 1-Target task), and b) change of gaze onto a displaced target with head free to move (2-Target task). 3) The 1-Target task requires the eyes to move in a direction opposite to that of the normal vestibulo-ocular reflex (VOR). The 2-Target task is identical, except that reorientation onto the new target calls for an initial saccadic eye movement in a direction opposite to that of the ensuing head movement, which is contrary to the normal pattern of eye-head coordination during gaze shifts. 4) Eye (EOG) and head (potentiometer) movements were continuously recorded (0-250 Hz) in an apparatus which permitted sudden, unexpected, electromagnetic braking of the head movement, either just before or during the intended manoeuvre. 5) Early adaptive strategies employed reduction of VOR gain, rearrangement of timing, amplitude and shape of "catch-up" saccades and the introduction of centrally programmed eye movements uncovered by the braking manoeuvres. 6) All of these phenomena were detectable in an initial series of 60 trials, in which the total exposure to visual-vestibular conflict was less than 30 s. They became more systematized and more marked after 6 h of active reversed vision experience. 7) Specifically, mean VOR gain, measured within the first 80 ms of head movement (deemed free of visuomotor influence), became markedly attenuated (25% in the first test series; 66% after 6 h of active vision-reversed exercise). In addition (not included in the above percentages) there were numerous occasions of complete absence of measurable VOR during head rotation, in both the first and final test series. 8) In the 1-Target task, the latency of the first "catch-up" saccade (re onset of head movement) tended to offset residual VOR by becoming shortened to the point of synchrony with head movement onset. This saccade (not present in control tests) continued to occur on those occasions when the head was unpredictably prevented from moving, and when head movements were made in the dark.(ABSTRACT TRUNCATED AT 400 WORDS)

Adaptation, Physiological

Eye-head coupling in humans. I. Simultaneous recording of isolated motor units in dorsal neck muscles and horizontal eye movements.

A tonic coupling between the horizontal component of eye position and dorsal neck muscle activity has been demonstrated in the cat and monkey. In order to demonstrate this synergy in humans and study its characteristics, we have measured the relation between the firing rate of individual motor unit of the splenius muscle and voluntary horizontal shifts of gaze using 5 degrees steps, in head-fixed subjects. Eye movement recording was achieved by conventional binocular electro-oculography and the activity of the right splenius muscle was recorded with Bronks coaxial bipolar electrodes inserted manually at the C4-C5 intervertebral level. The activity of 51 motor units in 10 subjects has been recorded. For all subjects, motor units firing rate increased when the gaze shifted to the ipsilateral side, and both increase in firing rate and recruitment were observed. These results demonstrate that the tonic eye head synergy is also present in man.

Action Potentials

Vertical eye movements related signals in antidromically identified medullary reticular formation neurons in the alert cat.

Short and long lead burst neurons antidromically activated from the rostral mesencephalic reticular formation, and synaptically activated from the contralateral superior colliculus were recorded in the medullary reticular formation underlying the prepositus hypoglossi nucleus. These neurons were shown to be related to vertical eye movements, ranging from pure vertical to oblique planes. Vertical saccade coding was similar to that of horizontal short lead pontine cells. The presence of vertical short and long lead burst neurons in the medullary reticular formation raises new questions about the organization of the control of eye movements in the vertical plane.

Action Potentials

Eye movements induced by off-vertical axis rotation (OVAR) at small angles of tilt.

Off-vertical rotation (OVAR) in darkness induced continuous horizontal nystagmus in humans at small tilts of the rotation axis (5 to 30 degrees). The horizontal slow eye velocity had two components: a mean velocity in the direction opposite to head rotation and a sinusoidal modulation around the mean. Mean velocity generally did not exceed 10 deg/s, and was less than or equal to the maximum velocity of optokinetic after-nystagmus (OKAN). Both the mean and modulation components of horizontal nystagmus increased with tilt angle and rotational velocity. Vertical slow eye velocity was also modulated sinusoidally, generally around zero. The amplitude of the vertical modulation increased with tilt angle, but not with rotational velocity. In addition to modulations in eye velocity, there were also modulations in horizontal and vertical eye positions. These would partially compensate for head position changes in the yaw and pitch planes during each cycle of OVAR. Modulations in vertical eye position were regular, increased with increases in tilt angle and were separated from eye velocity by 90 deg. These results are compatible with the interpretation that, during OVAR, mean slow velocity of horizontal nystagmus is produced by the velocity storage mechanism in the vestibular system. In addition, they indicate that the otolith organs induce compensatory eye position changes with regard to gravity for tilts in the pitch, yaw and probably also the roll planes. Such compensatory changes could be utilized to study the function of the otolith organs. A functional interpretation of these results is that nystagmus attempts to stabilize the image on the retina of one point of the surrounding world. Mean horizontal velocity would then be opposite to the estimate of head rotational velocity provided by the output of the velocity storage mechanism, as charged by an otolithic input during OVAR. In spite of the lack of actual translation, an estimate of head translational velocity could, in this condition, be constructed from the otolithic signal. The modulation in horizontal eye position would then be compensatory for the perceived head translation. Modulation of vertical eye velocity would compensate for actual changes in head orientation with respect to gravity.

Adult